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Systematics and evolution of the superfamily (Insecta: )

Dissertation

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

By

Huayan Chen, M.S.

Graduate Program in Entomology

The Ohio State University

2018

Dissertation Committee:

Norman F. Johnson, Advisor

Andy Michel

Hans Klompen

Mary M. Gardiner

Copyright by

Huayan Chen

2018

Abstract

The superfamily Platygastroidea (Insecta: Hymenoptera) is a diverse group of parasitic . Most species of Platygastroidea are egg of and spiders, or egg-larval parasitoids of gallflies, and many are recognized as important biological control agents of pests in agriculture and forestry. Further development of their potential use as biological agents depends upon a better understanding of the diversity and relationships within the superfamily.

The current limitations in phylogenetic knowledge have impeded research on these wasps. This dissertation has three primary goals: 1) document the diversity of three diverse genera,

Calliscelio, Habroteleia, and Chromoteleia; 2) reassess the evolutionary relationships within the superfamily Platygastroidea based on the analyses of four molecular markers from extensive taxon sampling; 3) characterize the molecular evolution of chemosensory receptor genes in the parasitic basalis.

Calliscelio is a cosmopolitan of parasitoids that attack the eggs of crickets

(: Gryllidae). In total, forty-two species are recognized from the New World, of which thirty-eight species are new to science. Four species are treated as junior synonyms of Calliscelio rubriclavus (Ashmead).

Habroteleia is found from Oriental region to Madagascar, Papua New Guinea, and the

Fijian archipelago. Seven species are recognized with four species are described as new to science. Four species are treated as junior synonyms of Habroteleia flavipes Kieffer. One species is treated as junior synonym of Habroteleia persimilis (Kozlov & Kononova). One species is transferred out of Habroteleia Kieffer. The status of two species is unclear but remains valid species.

i Chromoteleia species are mainly found from Neotrpoical region with the exception of one species from African continent. Twenty-seven species are recognized with twenty-one species are described as new. The identity and status of one species are unclear but it remains a valid species. All species are known only from the Neotropical region except for Chromoteleia congoana (Resbec), which only occurs in Africa.

To reassess the relationships within the superfamily Platygastroidea, Bayesian inference

(BI) and Maximum likelihood (ML) were conducted using four genes (18S, 28S, COI, and wingless), for 152 taxa representing 93 genera from all currently recognized platygastroid subfamilies and the majority of tribes. Both the ML and BI analyses of the concatenated data recovered similar well-supported clades. (Results). Implications for classification of the superfamily drawn from the phylogenetic relationships presented in this study are discussed.

Ancestral state reconstruction of host associations indicated that Orthoptera is the plesimorphic host group for the superfamily. Evolution of host shifts within the superfamily is also discussed.

To understand the potential role of chemosensory receptor genes in the host location and chemosensory speciation, the complete chemosensory receptor repertoires of T. basalis were identified via transcriptomic and genomic analyses. In total, 34 gustatory receptors (Grs), 170 odorant receptors (Ors) and one odorant co-receptor (Orco), and 23 inotropic receptors (Irs) were found in T. basalis. The phylogenetic relationships and expression profile of these chemosensory receptor genes were analyzed. The implications of these findings are also discussed.

ii Dedication

Dedicated to the sages who guide and lead me into the wonderful world of insects.

iii Acknowledgments First and foremost, I would like to thank my advisor, Dr. Norman Johnson, for accepting me as his student and supporting me all the way to this point. Dr. Johnson’s patience with my slow learning curve and mistakes, thoughtful advice both in research and teaching, and his persistent encouragement have been my vital source of motivation to become a better scientist. I would also like to thank the members of my advising committee, Dr. Andy Michel, Dr. Hans Klompen and Dr. Mary M. Gardiner for advising and supporting me throughout my study here at Ohio State. I have learned a lot from them both in the classroom and the lab. Thanks to Dr. John V. Freudenstein for helping me analyze data. A number of mentors have guided me to appreciate the beauties of insects and have eventually inspired me to pursue a career in the field of insect science. The late Dr. Zaifu Xu was the first mentor guided me into the field of Entomology. I am greatly indebted to him for his valuable guidance and continual support until his last days. I wish to express my gratitude to Dr. Lubomír Masner, who led me into the field of Platygastroidea. His endless enthusiasm in and philosophy in doing science have continually motivated me to move on. I would like to thank Dr. Kees van Achterberg, Dr. Junhua He, Dr. Massimo Olmi, and Dr. Giuseppe Fabrizio Turrisi for broadening my vision in Hymenoptera taxonomy. I would like to thank Dr. Elijah J. Talamas for the thoughtful discussions and collaboration we have had on Platyagastroidea. I am looking forward to more collaboration in the near future. I would also like to thank my officemate Zachary J. Lahey for assistance in analyzing data and of course for the friendships we have built. It is always enjoyable to discuss with him. Dr. Ovidiu A. Popovici is of helpful in developing the character matrix for Platyagastroidea and I appreciate his contribution. The three chapters of taxonomic work would not have been possible without the help of numerous taxonomists and curators of many insect collections around the world. I would like to thank the collectors of the 15,000 specimens I have studied. The molecular phylogeny of Platygastroidea could not have been done without the contributions of several members involved in the PBI project: Dr. Andy Austin, Dr. Andy Polaszek, Dr. Alejandro Valerio, and Dr. Charuwat Taekul. I am grateful to their help in providing specimens and collecting some of the molecular data.

iv Thanks to Dr. George Keeney for assistance in maintaining insect material for the transcriptomic study of . And, of course, thanks to the 10,000 wasps I bred, I appreciate their sacrifice in the discovery of chemosensory receptor genes. It has been enjoyable working in OSU insect collection because we have the most wonderful people here. I am particularly grateful to Dr. Luciana Musetti for always supporting me. I really appreciate her efforts in encouraging me to walk out my comfort zone and to enjoy the work and life here at Ohio State. I will definitely miss her. I owe immense thanks to Joseph Cora, who used to work here, for his assistance in conducting data analysis and using databases. I miss the days he accompanied me until 7 pm almost every workday for two years. I am also grateful to Sara Hemly for her help in entering specimen data, and making specimens available. I would like to thank my friends, Chundan Hong, Dr. Jingxian Liu, and Wei Dong, for their encouragement and support. I wish to thank my parents, Tai Chen and Ronmei Pan, my two younger brothers, Jingcai and Jingguo Chen, sister-in-law, Haiyun Yang, and my two lovely nieces, Qinying and Ziqing Chen. I could not have done it without their support and unconditional love. Lastly, I would like to express my gratitude to the below Chinese ancient verse which gives me hope whenever I feel desperate. 路漫漫其修远兮,吾将上下而求索 (The road ahead will be long and I’ll search up and down).

v Vita 2009...... B.S., South China Agricultural University, China 2012...... M.S., South China Agricultural University, China 2013–2014 ...... Graduate Research Associate, Department of Entomology, The Ohio State University 2015–2018 ...... Graduate Teaching Associate, Center for Life Science Education, The Ohio State University

Publications

Zheng B, Cao L, Tang P, van Achterberg C, Hoffmann AA, Chen H, Chen X, and Wei S (2018) Gene arrangement and sequence of mitochondrial genomes yield insights into the phylogeny and evolution of and sphecid wasps (Hymenoptera: ). Molecular Phylogenetics and Evolution 124: 1–9.

Chen H, Talamas EJ, Masner L, and Johnson NF (2018) Revision of the World species of the genus Habroteleia Kieffer (Hymenoptera, , ). ZooKeys 730: 85–120.

Zhu C, Luo S. Zhou X, Chen H, and Zhang G (2017) Earth BioGenome Project and Taxonomy. Science 25(11): 1251–1254. (Text in Chinese)

Chen H (2017) Biological collections: The Hidden Jewels. Science & Technology Review 35(18): 93. (Text in Chinese)

Chen H, Masner L, and Johnson NF (2017) New World species of the genus Calliscelio Ashmead (Hymenoptera, Platygastridae, Scelioninae). ZooKeys 648: 1–136.

Chen H, van Achterberg C, and Xu Z (2016) Descriptions of a new species of Foenatopus Smith from China and the male of Parastephanellus brevicoxalis (Hymenoptera, ). ZooKeys 612: 113–123.

Chen H, Hong C, van Achterberg C, and Xu Z (2016) Description of a new species of Pseudomegischus van Achterberg from China (Hymenoptera, Stephanidae). ZooKeys 601: 119– 125.

Chen H, Turrisi GF, and Xu Z (2016) A revision of the Chinese (Hymenoptera, ). ZooKeys 587: 77–124.

Chen H, van Achterberg C, He J, and Xu Z (2014) A revision of the Chinese Trigonalyidae (Hymenoptera, Trigonalyoidea). ZooKeys 385: 1–207.

Chen H, Johnson NF, Masner L, and Xu Z (2013) The genus Macroteleia Westwood

vi (Hymenoptera, Platygastridae, Scelioninae) from China. Zookeys 300: 1–98.

Chen H, Liu J, and Xu Z (2011) Description of a new platygastrid phenacocca (Hymenoptera) on Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae). Oriental Insects 45(2-3): 275–280.

Liu J, Chen H, and Xu Z (2011) Discovery of the female adult of genus Alareka Rajmohana et Narendran (Hymenoptera: ) from China. Oriental Insects 45(2-3): 181–185.

Liu J, Chen H, and Xu Z (2011) The genus Aneuropria Kieffer (Hymenoptera, Diapriidae), new record to China. Acta Zootaxonomica Sinica 36(4): 997–999.

Liu J, Chen H, and Xu Z (2011) Notes on the genus Haliday (Hymenoptera, Diapriidae) from China. ZooKeys 108: 49–60.

Liu J, Chen H, and Xu Z (2011) Two new species of the genus Foenobethylus Kieffer 1913 (Hymenoptera: ) from China with a key to the known species. Zootaxa 2806: 53–59.

Chen H, He L, Zheng C, Li P, Yi Q, and Xu Z (2011) Survey on the natural enemies of mealybug, Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae) from Guangdong and Hainan, China. Journal of Environmental Entomology 33(2): 269–272. (Text in Chinese)

Chen H, Cao R, and Xu Z (2010) First record of Aenasius bambawalei Hayat (Hymenoptera: ), a parasitoid of the mealybug Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae) from China. Journal of Enviornmental Entomology, 32(2): 280–282. (Text in Chinese)

Field of Study

Major Field: Entomology

vii Table of Contents Abstract ...... i Dedication ...... iii Acknowledgments...... iv Vita ...... vi List of Tables ...... x List of Figures ...... xi General Introduction ...... 1 Species Diversity and Taxonomy ...... 1 Phylogeny ...... 2 Chemosensory receptor genes in Trissolcus basalis...... 3 References ...... 6 Chapter 1: New World species of the genus Calliscelio Ashmead (Hymenoptera, Platygastridae, Scelioninae)...... 8 Abstract ...... 8 Introduction ...... 9 Materials and methods ...... 10 Taxonomy ...... 12 Key to separate Calliscelio, Calotelea, Holoteleia and Probaryconus ...... 18 Key to Females of Calliscelio of the New World ...... 20 Species descriptions ...... 29 Acknowledgments ...... 172 References ...... 172 Chapter 2: Revision of the World species of the Genus Habroteleia Kieffer (Hymenoptera, Platygastridae, Scelioninae) ...... 176 Abstract ...... 176 Introduction ...... 176 Materials and methods ...... 177 Taxonomy ...... 179 Key to separate Macroteleia, Triteleia and Habroteleia ...... 182 Key to Females ...... 182 Key to Males ...... 183 Species descriptions ...... 184 Acknowledgments ...... 217 References ...... 217 Chapter 3: Revision of the World species of the Genus Chromoteleia Ashmead (Hymenoptera, Platygastridae, Scelioninae) ...... 220 Abstract ...... 220 Introduction ...... 220 Materials and methods ...... 221 Taxonomy ...... 223 Key to Females ...... 228

viii Species descriptions ...... 235 Acknowledgments ...... 335 References ...... 335 Chapter 4: A molecular reassessment of the phylogeny of the superfamily Platygastroidea (Hymenoptera) ...... 338 Abstract ...... 338 Introduction ...... 338 Materials and Methods ...... 340 Results ...... 343 Discussion ...... 345 Acknowledgments ...... 364 References ...... 364 Chapter 5: Identification and expression analysis of chemosensory receptor genes in the egg parasitoid Trissolcus basalis ...... 369 Abstract ...... 369 Introduction ...... 369 Materials and Methods ...... 372 Results ...... 374 Discussion ...... 377 Acknowledgments ...... 400 References ...... 400 Bibliography ...... 404

ix List of Tables

Table 1.1 List of morphological terms and links to Hymenoptera Anatomy Ontology database for New World Calliscelio...... 170

Table 2.1 List of morphological terms and links to Hymenoptera Anatomy Ontology database for world species of Habroteleia...... 215

Table 3.1 List of morphological terms and links to Hymenoptera Anatomy Ontology database for New World Calliscelio...... 333

Table 4.1 Taxon sampling used in the molecular analysis listed in alphabetic order, including GenBank accession numbers. All GenBank accession numbers stating with MF and MH denote new sequences used for the first time in this study. The label data for all specimens have been geo‐referenced and recorded in the Hymenoptera On‐Line database. Details on the data associated with these specimens is publicly accessible at the following link, hol.osu.edu, and entering the specimen voucher ID in the form (e.g., OSUC 173847, note the space between the acronym and the number)...... 351

Table 4.2 Primers used to amplify four loci in this study...... 357

Table 4.3 Substitution models identified by PartitionFinder for each partition...... 358

Table 4.4 Summary statistics of genes used for phylogeny...... 358

Table 5.1 Grs identified from the genome and transcriptomes in T. basalis. Numbers in square brackets indicate partial exons. FA, female antenna; FB, female body; MA, male antenna; MB, male body...... 380

Table 5.2 Ors identified from the genome and transcriptomes in T. basalis. Numbers in square brackets indicate partial exons. FA, female antenna; FB, female body; MA, male antenna; MB, male body...... 382

Table 5.3 Irs identified from the genome and transcriptomes in T. basalis. FA, female antenna; FB, female body; MA, male antenna; MB, male body...... 391

x List of Figures

Figure 1.1 A, Calliscelio punctatifronssp. n., female, holotype (OSUC 191124), Antenna. B–C, Calliscelio punctatifronssp. n., male, paratype (OSUC 191125). B, Antenna; C, Tyloid. D, Calliscelio latifrons sp. n., female, holotype (OSUC 323077), Antenna. E–F, Calliscelio latifrons sp. n., male, paratype (OSUC 323075). E, Antenna; F, Tyloid.G, Calliscelio alcoa sp. n., female, holotype (OSUC 458212), Antenna. H–I, Calliscelio alcoa sp. n., male, paratype (OSUC 458222). H, Antenna; I, Tyloid.J, Calliscelio longius sp. n., female, paratype (OSUC 193935), Antenna. K–L, Calliscelio longius sp. n., male, paratype (OSUC 193596). K, Antenna; L, Tyloid...... 125

Figure 1.2 Fore wing, dorsal view. A, Calliscelio gatineau sp. n., female, holotype (OSUC 534340). B, Calliscelio storea sp. n., female, holotype (OSUC 546117). C, Calliscelio laticinctus Ashmead, female (OSUC 458242)...... 126

Figure 1.3 A, Calliscelio absconditumsp. n., female, paratype (OSUC 134791), dorsal habitus. B–F,Calliscelio absconditumsp. n., female, holotype (OSUC 323924). B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, , dorsal view...... 127

Figure 1.4 Calliscelio absum sp. n., female, holotype (OSUC 190716). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 128

Figure 1.5 Calliscelio alcoa sp. n., female, holotype (OSUC 458212). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 129

Figure 1.6 Calliscelio amadoi sp. n., female, holotype (OSUC 193589). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 130

Figure 1.7 Calliscelio armila sp. n., female, holotype (OSUC 458342). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 131

Figure 1.8 Calliscelio bidens sp. n., female, holotype (OSUC 458326). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 132

Figure 1.9 Calliscelio bisulcatus (Kieffer), female, holotype (CAS No. 9590). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 133

xi Figure 1.10 Calliscelio brachys sp. n., female, holotype (OSUC 193349). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 134

Figure 1.11 Calliscelio brevinotaulus sp. n., female, holotype (OSUC 190678). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 135

Figure 1.12 Calliscelio brevitas sp. n., female, holotype (OSUC 190705). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 136

Figure 1.13 Calliscelio carinigena sp. n., female, holotype (OSUC 343907). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 137

Figure 1.14 Calliscelio crater sp. n., female, holotype (OSUC 276803). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 138

Figure 1.15 Calliscelio crena sp. n., female, holotype (OSUC 553462). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 139

Figure 1.16 Calliscelio eboris sp. n., female, holotype (OSUC 193404). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 140

Figure 1.17 Calliscelio extenuatus sp. n., female, holotype (OSUC 48454). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 141

Figure 1.18 Calliscelio flavicauda sp. n., female, holotype (OSUC 553509). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 142

Figure 1.19 Calliscelio foveolatus sp. n., female, holotype (OSUC 532869). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 143

Figure 1.20 Calliscelio gatineau sp. n., female, holotype (OSUC 534340). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 144

xii Figure 1.21 Calliscelio glaber sp. n., female, holotype (OSUC 458136). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 145

Figure 1.22 Calliscelio granulatus sp. n., female, holotype (OSUC 534599). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 146

Figure 1.23 Calliscelio laticinctus Ashmead, female, syntype (USNMENT01059105). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Specimen labels...... 147

Figure 1.24 Calliscelio latifrons sp. n., female, holotype (OSUC 323077). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 148

Figure 1.25 Calliscelio levis sp. n., female, holotype (OSUC 553454). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 149

Figure 1.26 Calliscelio longius sp. n., female, holotype (OSUC 374730). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 150

Figure 1.27 Calliscelio magnificus sp. n., female, holotype (OSUC 458272). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 151

Figure 1.28 Calliscelio migma sp. n., female, holotype (OSUC 380181). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 152

Figure 1.29 Calliscelio minutia sp. n., female, holotype (OSUC 433847). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 153

Figure 1.30 Calliscelio paraglaber sp. n., female, holotype (OSUC 532684). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 154

Figure 1.31 Calliscelio pararemigio sp. n., female, holotype (OSUC 458239). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 155

xiii Figure 1.32 Calliscelio prolixus sp. n., female, holotype (OSUC 557587). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 156

Figure 1.33 Calliscelio punctatifrons sp. n., female, holotype (OSUC 191124). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 157

Figure 1.34 Calliscelio remigio sp. n., female, holotype (OSUC 458296). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 158

Figure 1.35 A–B, Acolus rubriclavus Ashmead, female, holotype (USNMENT00989042). A, Mesosoma and metasoma, lateral view; B, Mesosoma and metasoma, dorsal view. C–F, nigriceps Ashmead, male, syntype (USNMENT00989028). C, Mesosoma and metasoma, lateral view; D, Mesosoma, lateral view; E, Mesosoma and metasoma, dorsal view; F, Mesosoma, dorsal view...... 159

Figure 1.36 Caloteleia grenadensis Ashmead, female, holotype (B.M. TYPE HYM. 9.470). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Specimen labels...... 160

Figure 1.37 A–B, Caloteleia marlattii Ashmead, male, lectotype (USNMENT00989024). A, Lateral habitus; B, Dorsal habitus; C–F, Macroteleia ruskini Girault, female, holotype (USNMENT00989079). C, Mesosoma and metasoma, lateral view; D, Mesosoma and metasoma, dorsal view; E, Metasoma, dorsal view; F, Metasoma, ventral view...... 161

Figure 1.38 Calliscelio ruga sp. n., female, holotype (OSUC 553881). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 162

Figure 1.39 Calliscelio rugicoxa sp. n., female, holotype (OSUC 458325). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 163

Figure 1.40 Calliscelio sfina sp. n., female, holotype (OSUC 376920). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 164

Figure 1.41 Calliscelio storea sp. n., female, holotype (OSUC 546117). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 165

Figure 1.42 Calliscelio suni sp. n., female, holotype (OSUC 182752). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 166

xiv

Figure 1.43 Calliscelio telum sp. n., female, holotype (OSUC 276704). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 167

Figure 1.44 Calliscelio torqueo sp. n., female, holotype (OSUC 553598). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 168

Figure 1.45 Calliscelio virga sp. n., female, holotype (OSUC 534034). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F,Metasoma, dorsal view...... 169

Figure 2.1 A–B, Habroteleia mutabilis n. sp. A, Paratype (FBA 143219), Propodeum, dorsolateral view; B, Holotype (FBA 070892), Propodeum, dorsolateral view; C, Habroteleia salebra n. sp., male, paratype (OSUC 688063), Apex of metasoma, dorsal view; D, Habroteleia spinosa n. sp., male, paratype (OSUC 232878), Apex of metasoma, dorsal view; E, Habroteleia spinosa n. sp., female, holotype (OSUC 232889), Apex of metasoma, dorsal view; F, Habroteleia flavipes, male (OSUC 58007), Pronotum, lateral view...... 199

Figure 2.2 Baryconus vindhiensis, female, holotype (USNMENT01197073). A, Lateral habitus; B, Head, lateral view; C, Dorsal habitus; D, Labels...... 200

Figure 2.3 Habroteleia flavipes, female, holotype (MCSN 0001). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, lateral view; F, Metasoma and wings, dorsal view...... 201

Figure 2.4 Habroteleia bharatensis, female, holotype (USNMENT 01197132). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 202

Figure 2.5 Habroteleia browni, female, holotype (USNM Type No. 12894). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view...... 203

Figure 2.6 Chrestoteleia bakeri, female, holotype (MNHN 0013). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view...... 204

Figure 2.7 Habroteleia kotturensis, female, holotype (USNMENT01197074). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, lateral view; F, Metasoma and wings, dorsal view...... 205

xv Figure 2.8 Habroteleia mutabilis n. sp., female, holotype (FBA 142603). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view...... 206

Figure 2.9 Habroteleia mutabilis n. sp., female, paratype (FBA 070892). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view...... 207

Figure 2.10 Triteleia persimilis, female, holotype (ZMAS 0139). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 208

Figure 2.11 Triteleia dagavia, female, holotype (IEBR 0143). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Head and mesosoma, dorsal view; D, Propodeum, dorsolateral view; E, Head, lateral view; F, Metasoma, dorsal view...... 209

Figure 2.12 Phaedroteleia ruficoxa, male, holotype (MNHN_EY3427). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 210

Figure 2.13 Habroteleia salebra n. sp., female, holotype (OSUC 688076). A, Lateral habitus; B, Head and mesosoma, lateral view C, Dorsal habitus D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view...... 211

Figure 2.14 Habroteleia soa n. sp., female, holotype (CASENT 2136859). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head, anterior view; E, T5 and T6, dorsal view...... 212

Figure 2.15 Habroteleia spinosa n. sp., female, holotype (OSUC 232889). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view...... 213

Figure 2.16 A, Habroteleia flavipes, female (OSUC 688019), T5 and T6, dorsal view. B–C, Habroteleia persimilis, female (OSUC 687934). B, T5 and T6, dorsal view; C, Wings. .. 214

Figure 3.1 A–B, Chromoteleia tricarinata Kieffer, female (OSUC 268814). A, Antenna; B, Antennal clava, ventral view.C–D, Chromoteleia parvitas sp. n., holotype, female (OSUC 276182). C, Antenna; D, Antennal clava, ventral view; E–F, Chromoteleia longa sp. n., holotype, female (OSUC 320643). E, Antenna; F, Antennal clava, ventral view...... 291

Figure 3.2 A–B, Chromoteleia rara sp. n., female, holotype (OSUC 577495). A, Antenna; B, Antennal clava, ventral view.C, Chromoteleia ingens sp. n., paratype, male (OSUC 583458), Antenna; D, Chromoteleia bidens sp. n., paratype, male (OSUC 185675), Antenna...... 292

xvi Figure 3.3 Wings A, Chromoteleia longitarsis Kieffer, female (OSUC 584896). B, Chromoteleia robusta sp. n., female, paratype (OSUC 577466). C, Chromoteleia cuneus sp. n., female, holotype (OSUC 585001)...... 293

Figure 3.4 Mesopleuron and metapleuron. A, Chromoteleia connectens Kieffer, female, holotype (CAS TYPE9618). B, Chromoteleia longitarsis Kieffer, female (OSUC 584896)...... 294

Figure 3.5 A–B, Chromoteleia longitarsis Kieffer, female (OSUC 584896). A, Apical metasoma, dorsal view; B, Apical metasoma, lateral view. C–D Chromoteleia depilis sp. n., female, holotype (OSUC 577436). C, Apical metasoma, dorsal view; D, Apical metasoma, lateral view. E–F, Chromoteleia bidens sp. n., female, holotype (OSUC 577455). E, Apical metasoma, dorsal view; F, Apical metasoma, lateral view...... 295

Figure 3.6 A–B Chromoteleia fuscicornis Kieffer, female (OSUC 584322). A, Apical metasoma, dorsal view; B, Apical metasoma, lateral view. C, Chromoteleia pilus sp. n., male, paratype (OSUC 577941), Apical metasoma, dorsal view. D, Chromoteleia fuscicornis Kieffer, male (OSUC 586417), Apical metasoma, dorsal view. E, Chromoteleia bidens sp. n., male, paratype (OSUC 577455), Apical metasoma, dorsal view Apical. F, Chromoteleia longitarsis Kieffer, female (OSUC 584896), Hind tarsus. G, Chromoteleia depilis sp. n., female, paratype (OSUC 577436)), Hind tarsus. H, Chromoteleia bidens sp. n., female, paratype (OSUC 577455), Hind tarsus...... 296

Figure 3.7 Chromoteleia aequalis sp. n., female, holotype (OSUC 221942). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 297

Figure 3.8 Chromoteleia alternata sp. n., female, holotype (OSUC 136573). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 298

Figure 3.9 Chromoteleia bidens sp. n., female, holotype (OSUC 577455). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 299

Figure 3.10 Oxyscelio congoana Risbec, male, holotype (EY11376). A, Lateral habitus; B, Dorsal habitus; C, Head, anterior view; D, Head and mesosoma, dorsal view...... 300

Figure 3.11 Chromoteleia congoana (Risbec), female (OSUC 207829). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 301

Figure 3.12 Chromoteleia connectens Kieffer, female, holotype (CAS TYPE9618). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 302

xvii Figure 3.13 Chromoteleia brevitarsis Kieffer, female, holotype (CAS TYPE9597). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 303

Figure 3.14 Chromoteleia copiosa sp. n., female, holotype (OSUC 577518). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 304

Figure 3.15 Chromoteleia cuneus sp. n., female, holotype (OSUC 585001). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 305

Figure 3.16 Chromoteleia curta sp. n., female, holotype (OSUC 185626). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: light form) ...... 306

Figure 3.17 Chromoteleia curta sp. n., female, paratype (OSUC 149925). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: dark form) ...... 307

Figure 3.18 Chromoteleia depilis sp. n., female, holotype (OSUC 577436). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 308

Figure 3.19 Chromoteleia dispar sp. n., female, holotype (OSUC 190774). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 309

Figure 3.20 Chromoteleia feng sp. n., female, holotype (OSUC 233344). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 310

Figure 3.21 Chromoteleia fossa sp. n., female, holotype (OSUC 320582). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 311

Figure 3.22 Chromoteleia fuscicornis Kieffer, female, holotype (CAS TYPE9649). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head, anterior view...... 312

Figure 3.23 Chromoteleia fuscicornis Kieffer, female (OSUC 584322). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 313

xviii Figure 3.24 Chromoteleia ingens sp. n., female, holotype (OSUC 577500). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 314

Figure 3.25 Chromoteleia levitas sp. n., female, holotype (OSUC 577476). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 315

Figure 3.26 Chromoteleia longitarsis Kieffer, male, holotype (CAS TYPE9682). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 316

Figure 3.27 Chromoteleia longitarsis Kieffer, female (OSUC 584896). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 317

Figure 3.28 Chromoteleia longa sp. n., female, holotype (OSUC 188702). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 318

Figure 3.29 Chromoteleia maura sp. n., female, holotype (OSUC 584757). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 319

Figure 3.30 Chromoteleia parvitas sp. n., female, holotype (OSUC 586341). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 320

Figure 3.31 Chromoteleia pilus sp. n., female, holotype (OSUC 584261). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 321

Figure 3.32 Chromoteleia plana sp. n., female, holotype (OSUC 577460). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 322

Figure 3.33 Chromoteleia rara sp. n., female, holotype (OSUC 577495). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: small form) ..... 323

Figure 3.34 Chromoteleia rara sp. n., female, paratype (OSUC 557090). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: large form) ...... 324

xix Figure 3.35 Chromoteleia robusta sp. n., female, holotype (OSUC 577466). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 325

Figure 3.36 Chromoteleia semicyanea Ashmead, male, holotype (USNM Type No. 2251). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, female (OSUC 584765), Metasoma, dorsal view...... 326

Figure 3.37 Chromoteleia semilutea sp. n., female, holotype (OSUC 577506). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 327

Figure 3.38 Chromoteleia sparsa sp. n., female, holotype (OSUC 584751). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: light form) ...... 328

Figure 3.39 Chromoteleia sparsa sp. n., female, paratype (OSUC 583488). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: dark form) ...... 329

Figure 3.40 Chromoteleia tricarinata Kieffer, male, holotype (CAS TYPE9759). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 330

Figure 3.41 Chromoteleia tricarinata Kieffer, female (OSUC 584805). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view...... 331

Figure 3.42 Mesoscutum, mmc: median mesoscutal carina. A, Chromoteleia tricarinata Kieffer, male, holotype (CAS TYPE9759). B, Chromoteleia connectens Kieffer, female, holotype (CAS TYPE9618). C, Chromoteleia longitarsis Kieffer, male, holotype (CAS TYPE9682). D, Chromoteleia curta sp. n., female, holotype (OSUC 185626)...... 332

Figure 4.1 Net phylogenetic informativeness profiles for the four genes used in this study...... 359

Figure 4.2 Net phylogenetic informativeness profiles for the four genes and codon positions used in this study...... 360

Figure 4.3 Relationships of Platygastroidea from Bayesian analysis of concatenated 18S, 28S, COI and wingless sequences with 8 partitions. Posterior probabilities above 0.50 indicated on branches...... 361

xx Figure 4.4 Relationships of Platygastroidea from maximum likelihood analysis of concatenated 18S, 28S, COI and wingless sequences with 8 partitions. Boostrap values above 50 indicated on branches...... 362

Figure 4.5 Host associations plotted on the BI tree (see tree Fig. 4.3) using maximum parsimony method in Mesquite...... 363

Figure 5.1 Maximum likelihood phylogenetic tree of gustatory receptors (Grs) from T. basalis and other hymenopteran speices. Boostrap values above 50 indicated on branches. Am, A. mellifera, black; Mm, M. demolitor, orange; Nv, N. vitripennis, blue; Tb, T. basalis, red...... 393

Figure 5.2 Maximum likelihood phylogenetic tree of all odorant receptors (Ors) from T. basalis and representatives of Ors from other hymenopteran speices. Boostrap values above 50 indicated on branches. Am, A. mellifera, black; Mm, M. demolitor, orange; Nv, N. vitripennis, blue; Tb, T. basalis, red...... 394

Figure 5.3 Maximum likelihood phylogenetic tree of all odorant receptors (Ors) from T. basalis and other hymenopteran speices. Boostrap values above 50 indicated on branches. Am, A. mellifera, black; Mm, M. demolitor, orange; Nv, N. vitripennis, blue; Tb, T. basalis, red...... 395

Figure 5.4 Maximum likelihood phylogenetic tree of inotropic receptors (Irs) from T. basalis and other hymenopteran speices. Boostrap values indicated on branches. Am, A. mellifera, black; Mm, M. demolitor, orange; Nv, N. vitripennis, blue; Tb, T. basalis, red. 396

Figure 5.5 Tissue- and gender-specific expression profile of Grs in T. basalis. FA: female antenna; FB: female body; MA: male antenna; MB: male body...... 397

Figure 5.6 Tissue- and gender-specific expression profile of Ors in T. basalis. FA: female antenna; FB: female body; MA: male antenna; MB: male body...... 398

Figure 5.7 Tissue- and gender-specific expression profile of Irs in T. basalis. FA: female antenna; FB: female body; MA: male antenna; MB: male body...... 399

xxi General Introduction

Wasps classified as Platygastroidea (Hymenoptera) are one of the most diverse groups of egg parasitoids, comprising approximately 6000 described species (Aguiar et al. 2013; Johnson

2018). All platygastroids lay their own eggs in or on eggs or bodies of other insects and spiders.

Insect hosts include a wide range of nine orders (Austin et al. 2005), many of which are important pests in agriculture, forestry, and both human and health. These pests include locusts (Locusta migratoria, Choriocetes terminifera); stink bugs such as southern green stink bug (Nezara viridula), the brown marmorated stink bug (Halyomorpha halys Stål), and the sunn pest (Eurygaster integriceps); moths such as the fall armyworm (Spodoptera frugiperda) and gypsy moth (Lymantria dispar), Hessian (Mayetiola destructor); and mealy bugs

(Phenacoccus solenopsis, Oracella acuta) (Austin et al. 2005). Many platygastroid species are considered to be important biological control agents of insect pests and a number of them have been deployed successfully for that purpose (Austin et al. 2005; Taekul et al. 2014).

In addition, due to their taxonomical and biological diversity, platygastroids also serve as ideal systems in many areas of entomological research, particularly in studying the evolution of parasitoid-host relationships (Austin et al. 1997, 2005; Murphy et al. 2007; Taekul et al. 2014).

Despite their scientific and economic importance, Platygastroidea remains a large void in our knowledge of the tree of life.

Species Diversity and Taxonomy

Currently, Platygastroidea is comprised of five subfamilies: ,

Sceliotrachelinae, Scelioninae, , and . The subfamily Scelioninae is particularly species-rich and diverse in host associations. This subfamily contains many diverse

1 genera. Recent revisions on some of these genera, e.g., Oxyscelio (Burks et al. 2013a, 2013b,

2016), Macroteleia (Chen et al. 2013), and Scelio (Yoder et al. 2014), etc., have demonstrated that many new species still need to be described. Taxonomic revisions are essential for the fundamental understanding of biodiversity and its conservation. In order to understand the diversity of these beneficial wasps, one of the objectives of this dissertation is to describe and delimit the New World species of the genus Calliscelio (Chapter 1), and the world species of the genera Habroteleia (Chapter 2) and Chromoteleia (Chapter 3). These taxonomic revisions have significantly advanced out understanding of the diversity and biogeography of these parasitoids, amongst which there might be some that will eventually prove to be useful in pest management.

Phylogeny

The number of higher-level groups and species combined with their rapid radiation has previously impeded efforts to meaningfully sample across the major lineages for resolving the evolutionary history of Platygastroidea (Austin et al. 2005). The lack of a robust hypothesis of evolutionary relationships has impeded research on these wasps such as taxonomic classification, dynamics of speciation and , evolution of phenotypic, behavioral, and many other biological traits (Austin et al. 2005). Higher-level evolutionary relationships within

Platygastroidea remain unclear (Murphy et al. 2007; Taekul et al. 2014) and the current classification does not reflect its trure evolutionary history. In addition, host shift patterns are crucial to understanding the evolution and mechanisms of host finding strategies among these parasitoids. Such patterns cannot be inferred without a robust phylogeny for the superfamily.

A major difficulty in resolving the relationships among platygastroids is due to the lack of informative morphological characters. Molecular data would seem to be useful in resolving

2 this issue. Murphy et al. (2007) provided the first most extensive analysis of platygastroids based on molecular data from 18S rRNA, 28S rRNA and CO1 genes. However, their results indicated that none of the currently-recognized tribes are monophyletic. In addition, their conclusion that the subfamily Telenominae was monophyletic was disputed by a recent molecular study (Taekul et al. 2014).

Although the data from Murphy et al. (2007) are much more extensive than any other available data set, molecular data for all three genes studied were available for only 46 of the 70 ingroup taxa. New studies including more taxa and additional gene markers should provide a sharper picture of the phylogenetic relationships within the family.

To this end, Chapter 4 addresses phylogenetic and taxonomic hypotheses across multiple hierarchical levels within the superfamily based on the phylogenetic analyses of four molecular markers (18S, 28S, COI, and wingless) from 152 ingroup taxa.

Chemosensory receptor genes in Trissolcus basalis

Like other parasitic wasps, to successfully suppress their hosts and reproduce themselves, platygastroids rely on their ability of chemosensation to locate food, mates, hosts and other resources in a complex environment. These chemosensory activities are conducted by a range of proteins housed in the olfactory sensory neurons (OSNs) of different types of chemosensory sensilla that are located on peripheral appendages, such as the legs, wings, maxillary and labial palps, and especially the antennae (Suh et al. 2014; McKenzie et al. 2016). These proteins are encoded in moderately sized multigene families, including genes coding for the odorant-binding proteins (OBPs), chemosensory proteins (CSPs), sensory neuron membrane proteins (SNMPs) and a chemoreceptor superfamily formed by the gustatory receptors (GRs), odorant receptors

3 (ORs), and ionotropic receptors (IRs) (Zhou et al. 2015). Different chemoreceptors can recognize specific chemical cues.

The molecular features of chemosensory genes can help to unravel the mechanisms of chemical perception from molecules to behavior in insects (Smadja and Butlin 2009). Therefore, an evolutionary comparison of these genes from different groups of insects may shed light on the chemosensory mechanisms of host adaptation and speciation in diverse insect taxa (Hallem et al.

2006). For example, chemosensory receptor genes are one of the fastest evolving gene families in insects (Neafsey et al. 2015) and the expansion in one subfamily of odorant receptors is believed to play an important role in the evolution of sociality in honey bees (Robertson &

Wanner 2006) and (Zhou et al. 2012; Engsontia et al. 2015; McKenzie et al. 2016). In addition, functional characterization of chemosensory genes may prove useful in the design of novel approaches to manipulate the behaviors of economically and medically important insects

(Suh et al 2014).

The parasitic wasp Trissolcus basalis (Hymenoptera: Platygastridae) is an ideal model for investigating the evolution of chemosensory receptor genes. First, it is an endoparasitoid that attacks the eggs of stinkbugs and has been widely used as an effective biological control agent of the pan-tropical agricultural pest Nezara viridula (Hemiptera: Pentatomidae) in many countries

(Loch & Walter 1999). This wasp uses multiple chemical cues to find mates (Loch & Walter

2002) and hosts (Mattiacci et al. 1993; Rosi et al. 2001; Colazza et al. 2009). In addition, the genus Trissolcus contains both generalists and specialists and has 193 described species. The identification of chemosensory receptor genes in T. basalis would help us understand the underlying mechanisms controlling chemical communication, behavior, host selection and acceptance, and host-wasp interactions so that we can design better biological control strategies

4 and develop monitoring tools to track parasitism rates in the field both for this species and related taxa. Comparing the evolutionary patterns of chemosensory receptors among different groups of parasitic wasps may allow us to understand the underlying mechanisms of chemosensory speciation. Chapter 5 identifies the chemosensory receptor gene repertoires and characterizes their molecular evolution in T. basalis via a genome wide survey and transcriptomic analysis.

5 References Aguiar AP, Deans AR, Engel MS, Forshage M, Huber JT, Jennings JT, Johnson NF, Lelej AS, Longino JT, Lohrmann V, Mikó I, Ohl M, Rasmussen C, Taeger A, Yu DSK (2013) Order Hymenoptera. Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa 3703: 51–62. Austin AD, Field SA (1997) The ovipositor system of scelionid and platygastrid wasps (Hymenoptera: Platygastroidea): comparative morphology and phylogenetic implications. Invertebrate Taxonomy 11: 1–87. Austin AD, Johnson NF, Dowton M (2005) Systematics, evolution, and biology of scelionid and platygastrid wasps. Annual Review of Entomology 50: 553–582. Burks RA, Masner L, Johnson NF, Austin AD (2013a) Systematics of the parasitic wasp genus Oxyscelio Kieffer (Hymenoptera, Platygastridae s.l.), part I: Indo-Malayan and Palearctic fauna. ZooKeys 292:1−263. Burks RA, Masner L, Johnson NF, Austin AD (2013b) Systematics of the parasitic wasp genus Oxyscelio Kieffer (Hymenoptera, Platygastridae s.l.), part II: the Australian and southwest Pacific fauna. ZooKeys 331: 1−266. Burks RA, Masner L, Johnson NF, Austin AD (2016) Systematics of the parasitic wasp genus Oxyscelio Kieffer (Hymenoptera, Platygastridae s.l.), part III: African fauna. ZooKeys 565: 29–71. Chen H, Johnson NF, Masner L, Xu Z (2013) The genus Macroteleia Westwood (Hymenoptera, Platygastridae, Scelioninae) from China. Zookeys 300: 1−98. Colazza S, Lo Bue M, Lo Giudice D, Peri E (2009) The response of Trissolcus basalis to footprint contact kairomones from Nezara viridula females is mediated by leaf epicuticular waxes. Naturwissenschaften 96: 975−981. Engsontia P, Sangket U, Robertson HM, Satasook C (2015) Diversification of the odorant receptor gene family and positive selection on candidate cuticular hydrocarbon receptors. BMC Research Notes 8: 380. Hallem EA, Dahanukar A, Carlson JR (2006) Insect odor and taste receptors. Annual Review of Entomology 51: 113–135. Johnson NF (2018) Platygastroidea Online. http://osuc.biosci.ohio- state.edu/hymDB/eol_scelionidae.home Loch AD, Walter GH (1999) Multiple host use by the egg parasitoid Trissolcus basalis (Wollaston) in a soybean agriculture system: biological control and environmental implications. Agricultural and Forest Entomology 1: 271–280. Loch AD, Walter GH (2002) Mating behavior of Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae): potential for outbreeding in a predominantly inbreeding species. Journal of Insect Behavior 15: 13−23. Mattiacci L, Vinson SB, Williams HJ, Aldrich JR, Bin F (1993) A long range attractant kairomone for egg parasitoid Trissolcus basalis, isolated from defensive secretion of its host, Nezara viridula. Journal of Chemical Ecology 19: 1167−1181. McKenzie SK, Fetter-Pruneda I, Ruta V, Kronauer DJC (2016) Transcriptomics and neuroanatomy of the clonal raider ant implicate an expanded clade of odorant receptors in chemical communication. Murphy NP, Carey D, Castro LR, Dowton M, Austin AD (2007) Phylogeny of the platygastroid wasps (Hymenoptera) based on sequences from the 18S rRNA, 28S rRNA and cytochrome

6 oxidase I genes: implications for the evolution of the ovipositor system and host relationships. Biological Journal of the Linnean Society 91: 653–669. Neafsey DE, et al. (2015) Mosquito genomics. Highly evolvable malaria vectors: the genomes of 16 Anopheles mosquitoes. Science 347:1258522. Robertson HM, Wanner KW (2006) The chemoreceptor superfamily in the honeybee Apis mellifera: expansion of the odorant, but not gustatory, receptor family. Genome Research 16:1395–1403. Rosi MC, Isidoro N, Colazza S, Bin F (2001) Source of the host marking pheromone in the egg parasitoid Trissolcus basalis (Hymenoptera: Scelionidae). Journal of Insect Physiology 47: 989–995. Smadja C, Butlin RK (2009) On the scent of speciation: the chemosensory system and its role in premating isolation. Heredity 102: 77−97. Suh E, Bohbot JD, Zwiebel LJ (2014) Peripheral olfactory signaling in insects. Current Opinion in Insect Science 6: 86−92. Taekul C, Valerio AA, Austin AD, Klompen H, Johnson NF (2014) Molecular phylogeny of telenomine egg parasitoids (Hymenoptera: Platygastridae s.l.: Telenominae): evolution of host shifts and implications for classification. Systematic Entomology 39: 24–35. Yoder MJ, Valerio AA, Polaszek A, van Noort S, Masner L, Johnson NF (2014) Monograph of the Afrotropical species of Scelio Latreille (Hymenoptera: Platygastridae), egg parasitoids of acridid (Orthoptera: ). ZooKeys 380: 1−188. Zhou X, Slone JD, Rokas A, Berger SL, Liebig J, Ray A, Reinberg D, Zwiebel LJ (2012) Phylogenetic and transcriptomic analysis of chemosensory receptors in a pair of divergent ant species reveals sex-specific signatures of odor coding. PLoS Genet 8: e1002930. Zhou X, Rokas A, Berger SL, Liebig J, Ray A, Zwiebel LJ (2015) Chemoreceptor evolution in hymenoptera and its implications for the evolution of eusociality. Genome Biology and Evolution 7: 2407–2416.

7 Chapter 1: New World species of the genus Calliscelio Ashmead (Hymenoptera, Platygastridae, Scelioninae)

Abstract The genus Calliscelio Ashmead is presumed to be a diverse group of parasitoids of the eggs of crickets (Orthoptera: Gryllidae). A least one species has been found to be an important factor in depressing cricket pest populations. The New World species of Calliscelio are revised. Forty-two species are recognized, 3 are redescribed: C.bisulcatus (Kieffer), C. laticinctus Ashmead, C. rubriclavus (Ashmead), comb. n.; and 38 are described as new: C. absconditum Chen & Johnson, sp. n., C. absum Chen & Johnson, sp. n., C. alcoa Chen & Masner, sp. n., C. amadoi Chen & Johnson, sp. n., C. armila Chen & Masner, sp. n., C. bidens Chen & Masner, sp. n., C. brachys Chen & Johnson, sp. n., C. brevinotaulus Chen & Johnson, sp. n., C. brevitas Chen & Johnson, sp. n., C. carinigena Chen & Johnson, sp. n., C. crater Chen & Johnson, sp. n., C. crena Chen & Johnson, sp. n., C. eboris Chen & Johnson, sp. n., C. extenuatus Chen & Johnson, sp. n., C. flavicauda Chen & Johnson, sp. n., C. foveolatus Chen & Johnson, sp. n., C. gatineau Chen & Johnson, sp. n., C. glaber Chen & Masner, sp. n., C. granulatus Chen & Masner, sp. n., C. latifrons Chen & Johnson, sp. n., C. levis Chen & Johnson, sp. n., C. longius Chen & Johnson, sp. n., C. magnificus Chen & Masner, sp. n., C. migma Chen & Johnson, sp. n., C. minutia Chen & Johnson, sp. n., C. paraglaber Chen & Johnson, sp. n., C. pararemigio Chen & Masner, sp. n., C. prolixus Chen & Johnson, sp. n., C. punctatifrons Chen & Johnson, sp. n., C. remigio Chen & Masner, sp. n., C. ruga Chen & Johnson, sp. n., C. rugicoxa Chen & Masner, sp. n., C. sfina Chen & Johnson, sp. n., C. storea Chen & Johnson, sp. n., C. suni Chen & Johnson, sp. n., C. telum Chen & Johnson, sp. n., C. torqueo Chen & Johnson, sp. n., C. virga Chen & Johnson, sp. n. Four species are treated as junior synonyms of Calliscelio rubriclavus (Ashmead): Anteris nigriceps Ashmead syn. n., Caloteleia marlattii Ashmead syn. n., Caloteleia grenadensis Ashmead syn. n., and Macroteleia ruskini Girault syn. n.

8 Introduction

The genus Calliscelio Ashmead was first erected for a single species, C. laticinctus Ashmead, from the Lesser Antilles (Ashmead, 1893). For nearly 80 years it was considered to be a valid genus although it remained narrowly defined. Kieffer (1926), for example, included only two species in his monograph on world Scelionidae (= Platygastridae sensu Sharkey 2007). Masner (1972) treated Calliscelio as a junior synonym of Calotelea Westwood. Shortly thereafter, though, Masner (1976) rejected this hypothesis, treated Calliscelio as valid, and also significantly expanded the extent of the genus. Kieffer had described seven other genera over the years, all of which Masner (1976) treated as junior synonyms of Calliscelio (see generic synonymy below). Recently, in a survey of external morphology across the superfamily Platygastroidea, Talamas et al. (2016) found that the prioniform sensillum on the mandible is synapomorphy for the genus Calliscelio and treated Crama Galloway, Lispoteleia Galloway, Xentor Masner and Johnson, and Yunkara Galloway as junior synonyms of Calliscelio. The current concept of Calliscelio is a relatively large genus in the subfamily Scelioninae, comprising 77 known species (including 3 fossil species; Johnson 2016). It is a worldwide genus, with species found in all six major biogeographic regions. Interestingly, at least two species, i.e., C. rubriclavus (Ashmead) and C. elegans (Perkins), have extraordinarily broad geographic distributions possibly due to increased globalization of human commerce (Masner et al., 2009). This phenomenon leads to the question: from which part of the world did these species originate? Also, although Calliscelio was originally erected based on species in the New World, only 6 species were reported from this region to date, suggesting that more species remain to be discovered. Thus, a comprehensive revision of species of Calliscelio based on extensive sampling from the New World is needed to provide a more accurate understanding of the diversity and interrelationships among species in the genus. Even though Calliscelio is a diverse, common, and widespread genus, the available biological data are extremely scanty. The only host record is for C. teleogrylli Hill, which was reported to attack the eggs of Teleogryllus commodus (Walker) (Orthoptera: Gryllidae), a widespread pest in pastures in the Pacific region (Hill, 1983). By comparison with C. teleogrylli, Masner et al. (2009) suggested that C. elegans might attack the eggs of one or more species of Gryllidae associated with sugar cane.

9 The goal of this work is to reevaluate the known species of Calliscelio from the New World, expand the biogeographic data associated with these species, and to describe new species. The contributions of the authors are as follows. H-y Chen and N.F. Johnson: character definition, generic concept development, species concept development, imaging, key development, manuscript preparation; L. Masner: character definition, generic concept development, species concept development. The authors of the new species are indicated in the heading of each description.

Materials and methods

This work is based upon specimens in the following collections, with abbreviations used in the text: 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; CAS, California Academy of Sciences, San Francisco, CA; CMNH, Carnegie Museum of Natural History, Pittsburgh, PA; CNCI, Canadian National Collection of Insects, Ottawa, Canada; FSCA, Florida State Collection of , Gainesville, FL; IAVH, Instituto Alexander von Humboldt, Villa de Leyva, Columbia; MNHNPY, Museo Nacional de Historia Natural del Paraguay, San Lorenzo; INHS, Illinois Natural History Survey, Urbana, IL; MAIC, M.A. Ivie Private Collection, Bozeman, MT; MEMU, Mississippi State University, Mississippi State, MS; MHNG, Muséum d'Histoire Naturelle, Geneva, Switzerland; MNHN, Muséum National d'Histoire Naturelle, Paris, France; MPEG, Museu Paraense Emílio Goeldi, Belém, PA, Brazil; MZLU, Lund Museum of Zoology, Lund University, Lund, Sweden; MZSP, Museu de Zoologia da Universidade de São Paulo Sao, Brazil; NMNH, National Museum of Natural History, Washington, DC; OSUC, C.A. Triplehorn Insect Collection, Ohio State University, Columbus, OH; UCDC, R.M. Bohart Museum of Entomology, University of California, Davis, CA; UCFC, The Stuart M. Fullerton Collection of Arthropods at the University of Central Florida, Orlando, FL; UCMC, University of Colorado, Boulder, CO;UCRC, University of California, Riverside, CA; USNM, National Museum of Natural History, Washington, DC. 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

10 antennomeres of the female, with the segment interval specified followed by the number of sensilla per segment (Bin, 1981); EH: eye height, length of compound eye measured parallel to dorsoventral midline of head; IOS: interocular space, minimal distance on frons between compound eyes; OD: ocellar diameter, greatest width of ocellus; OOL: ocular ocellar line, shortest distance from inner orbit and outer margin of posterior ocellus (Masner 1980); T1, T2, ... T7: metasomal tergite 1, 2, ... 7; S1, S2, … S7: metasomal sternite 1, 2, … 7. Morphological terminology otherwise generally follows Masner (1980) and Mikó et al. (2007). Morphological terms used in this work are as in the Hymenoptera Anatomy Ontology (Yoder et al. 2010) (Table 1.1). Identifiers (URIs) in the format HAO_XXXXXXX represent concepts in the HAO and are provided to enable readers to confirm their understanding of the concepts being referenced. To find out more about a given concept, including additional images, notes, references and other metadata, use the identifier as a search term at http://glossary.hymao.org or use the identifier as a web-link. In the Material Examined section, the specimens studied are recorded in an abbreviated format, using unique identifiers (numbers prefixed with “OSUC”, “CASENT”, “UCFC”, “CMNH”, “MHNG”) for the individual specimens. The label data for all specimens have been georeferenced and recorded in the Hymenoptera On-Line database, and details on the data associated with these specimens can be accessed at the following link, hol.osu.edu, and entering the identifier in the form (note the space between the acronym and the number). 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, www.zoobank.org), and other taxonomic names, where appropriate, have been retrospectively registered. Data associated with the genus Calliscelio can be accessed at hol.osu.edu/index.html?id=461. The generic and species descriptions were generated by an xBio:D platform application, vSysLab (vsyslab.osu.edu), designed to facilitate the production of a taxon by character data matrices, and to integrate those data with the existing taxonomic, bibliographic 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:

11 Character state (s)". Polymorphic characters are indicated by semicolon-separated character states. Images and measurements were made using Combine ZP and AutoMontage extended-focus software, using JVC KY-F75U digital camera, Leica Z16 APOA microscope, and 1X objective lens. Images were post-processed with Adobe Photoshop CS3 Extended. A standard set of images is provided for each species: dorsal habitus, lateral habitus, dorsal and lateral views of the head and mesosoma, and anterior view of head. The individual images are archived in Specimage (specimage.osu.edu), the image database at The Ohio State University.

Taxonomy

Calliscelio Ashmead

Calliscelio Ashmead, 1893: 209, 218 (original description. Type: Calliscelio laticinctus Ashmead, by monotypy and original designation, keyed); Ashmead, 1894: 216 (keyed); Dalla Torre, 1898: 501 (catalog of species); Ashmead, 1900: 327 (list of species of West Indies); Ashmead, 1903: 91 (keyed); Brues, 1908: 27, 28, 33 (diagnosis, list of species, keyed); Kieffer, 1908: 122 (keyed); Kieffer, 1910b: 66 (keyed); Kieffer, 1913: 232 (description); Kieffer, 1926: 273, 499 (description, keyed, key to species); Muesebeck & Walkley, 1956: 338 (citation of type species); Baltazar, 1966: 185 (cataloged, catalog of species of the Philippines); Masner, 1972: 839 (junior synonym of Calotelea Westwood); Masner, 1976: 34, 36, 43 (description; key to Calliscelio Ashmead, Paridris Kieffer, Oethecoctonus Ashmead, and Probaryconus Kieffer; key to Calotelea Westwood and Calliscelio Ashmead); Mani & Sharma, 1982: 178 (description); Galloway & Austin, 1984: 8, 27, 28 (description, list of species described from Australia, keyed); Kozlov & Kononova, 1985: 19 (description, key to species of the Palearctic); Kozlov & Kononova, 1990: 19, 173, 183 (description, key to species of the USSR, keyed); Johnson, 1992: 355 (catalog of world species); Kononova, 1995: 61, 69 (keyed, diagnosis, key to species of Russian Far East); Austin & Field, 1997: 20, 68 (structure of ovipositor system, discussion of phylogenetic relationships); Narendran & Ramesh Babu, 1990: 2 (key to species of India); Lê, 2000: 31, 46 (keyed, description, key to species); Loiácono & Margaría, 2002:

12 557 (catalog of Brazilian species); Mineo, 2004: 174 (distribution in Sicily); Rajmohana, 2006: 116, 119, 120 (description, keyed, key to species of India); Kononova & Fursov, 2007a: 57 (description); Kononova & Fursov, 2007b: 98 (description); Kononova & Kozlov, 2008: 23, 257, 258 (description, keyed, key to species of Palearctic region); Rajmohana & Peter, 2013: 76 (key to species Calliscelio rugosus Rajmohana & Peter and Calliscelio agaliensis Narendran & Ramesh Babu); Talamas & Buffington, 2015: 12. (fossil in Dominican ); Talamas, Johnston-Jordan & Buffington, 2016: 413, 416 (description, synonymy). http://zoobank.org/29B1D7E4-1173-4D61-B695-CA755632F5EA http://bioguid.osu.edu/xbiod_concepts/461 Baryteleia Kiefffer, 1926: 273, 544 (original description. Type: Macroteleia nigriceps Kieffer, by original designation, keyed, key to species); Muesebeck & Walkley, 1956: 336 (citation of type species); Masner, 1976: 36 (junior synonym of Calliscelio Ashmead). http://zoobank.org/AB0DBC82-18D8-431F-9069-783F4874CEC1 http://bioguid.osu.edu/xbiod_concepts/8406 Caenoteleia Kieffer, 1926: 266, 550 (original description. Type: Caloteleia elegans Perkins, by monotypy, keyed); Muesebeck & Walkley, 1956: 338 (citation of type species); Johnson, 1992: 355 (catalog of world species); Masner et al., 2009: 60 (junior synonym of Calliscelio Ashmead, discussion of status). http://zoobank.org/5FEFDDD1-26AD-40D7-A498-BB4B5DF03630 http://bioguid.osu.edu/xbiod_concepts/460 Ceratoteleia Kieffer, 1908: 121 (original description. TypeCaloteleia grenadensis Ashmead, designated by Kieffer (1926), keyed); Kieffer, 1910b: 65, 66, 88 (description, list of species, keyed); Dodd, 1913a: 131, 144 (key to species of Australia); Dodd, 1913b: 176 (comparison with Macroteleia Westwood); Kieffer, 1913: 222 (description); Kieffer, 1913: 232 (description); Kieffer, 1914a: 315 (description, key to species of Europe and Algeria); Kieffer, 1926: 273, 500 (description, keyed, key to species, designation of type species); Nixon, 1931: 356 (keyed, key to species of Africa); Nixon, 1933: 292 (keyed); Brues, 1940: 82 (key to species of Baltic amber); Maneval, 1940: 114 (keyed); Risbec, 1950: 603 (key to species of Ethiopian region); Muesebeck & Walkley, 1951: 705 (catalog of species of U.S.

13 and Canada); Muesebeck & Walkley, 1956: 341 (citation of type species); Masner, 1976: 36 (junior synonym of Calliscelio Ashmead). http://zoobank.org/70CBF446-7CD7-4211-BF0C-F401DAE92DDA http://bioguid.osu.edu/xbiod_concepts/8400 Crama Galloway, 1984: 7, 8, 28 (original description. Type: Baryconus albicoxa Dodd, by original designation, key to Australian species, keyed); Johnson, 1992: 364 (catalog of world species); Talamas, Johnston-Jordan & Buffington, 2016: 413, 417 (junior synonym of Calliscelio Ashmead). http://zoobank.org/10BDF90E-0D3E-491E-98B2-8B39BB9D5871 http://bioguid.osu.edu/xbiod_concepts/466 Glyptoteleia Kieffer, 1926: 272, 487 (original description. Type: Baryconus bisulcatus Kieffer, by monotypy, keyed); Muesebeck & Walkley, 1956: 356 (citation of type species); Szabó, 1962: 241 (diagnosis); Masner, 1976: 36 (junior synonym of Calliscelio Ashmead); De Santis, 1980: 312 (catalog of species of Brazil). http://zoobank.org/FFA0AF0B-A126-4A36-8DB4-E25910A88C3D http://bioguid.osu.edu/xbiod_concepts/8405 Lispoteleia Galloway, 1984: 7, 9, 35 (original description. Type: Lispoteleia collina Galloway, by original designation, key to species of Australia, keyed); Johnson, 1992: 421 (catalog of world species); Austin and Field, 1997: 22, 68 (structure of ovipositor system, discussion of phylogenetic relationships); Talamas, Johnston-Jordan & Buffington, 2016: 413, 417 (junior synonym of Calliscelio Ashmead). http://zoobank.org/A458DE09-DAFA-424E-B60E-1A831B19D01E http://bioguid.osu.edu/xbiod_concepts/503 Mesoteleia Kieffer, 1917: 51 (original description. Type: Mesoteleia pallida Kieffer, by monotypy and original designation); Kieffer, 1926: 271, 441 (description, keyed); Muesebeck & Walkley, 1956: 369 (citation of type species); Baltazar, 1966: 182 (cataloged, catalog of species of the Philippines); Masner, 1976: 36 (junior synonym of Calliscelio Ashmead). http://zoobank.org/B6CD6365-672B-4E49-B3AE-6AAFA9C42DA6 http://bioguid.osu.edu/xbiod_concepts/8404

14 Prosanteris Kieffer, 1908: 121, 136 (original description. Type: Anteris nigriceps Ashmead, designated by Kieffer (1910b), keyed); Kieffer, 1910b: 65, 87 (description, key to subgenera, list of species, keyed); Kieffer, 1913: 232 (description); Kieffer, 1926: 272, 437 (description, keyed, key to species); Muesebeck & Walkley, 1951: 704 (catalog of species of U.S. and Canada); Muesebeck & Walkley, 1956: 391 (citation of type species); Muesebeck, 1958: 93 (junior synonym of Ceratoteleia Kieffer). http://zoobank.org/31DB13FB-2699-4E1D-A023-1BA21C1B9503 http://bioguid.osu.edu/xbiod_concepts/8401 Uroscelio Kieffer, 1914: 291 (original description. Type: Uroscelio luteipes Kieffer, by monotypy and original designation); Kieffer, 1926: 268, 409 (description, keyed); Muesebeck & Walkley, 1956: 408 (citation of type species); Baltazar, 1966: 180 (cataloged, catalog of species of the Philippines); Masner, 1976: 36 (junior synonym of Calliscelio Ashmead). http://zoobank.org/91A2A4C7-7D11-4723-8C53-02DEA8A0213B http://bioguid.osu.edu/xbiod_concepts/8403 Xentor Masner and Johnson, 2007: 12, 14 (original description. Type: Xentor schlingeri Masner and Johnson, by original designation, key to species); Talamas, Johnston-Jordan & Buffington, 2016: 416 (junior synonym of Calliscelio Ashmead). http://zoobank.org/1578C9FB-4A24-42D6-922D-05CDA2626906 http://bioguid.osu.edu/xbiod_concepts/211604 Yunkara Galloway, 1984: 9, 33 (original description. Type: Yunkara inornata Galloway, by monotypy and original designation, keyed); Johnson, 1992: 510 (catalog of world species); Talamas, Johnston-Jordan & Buffington, 2016: 413, 418 (junior synonym of Calliscelio Ashmead). http://zoobank.org/4FBE9CB9-3B71-4DFB-BCE0-781664A31929 http://bioguid.osu.edu/xbiod_concepts/578

Description (based on New World species). Length: 1.27–3.88 mm; body moderately to markedly elongate, robust. Head. Head shape in dorsal view: transverse. Hyperoccipital carina: absent; present. Occipital carina: present, complete medially; present laterally, broadly interrupted medially;

15 completely absent. Occipital carina sculpture: crenulate; unsculptured. OOL: lateral ocellus nearly contiguous with inner orbits, OOL < 0.5 OD; lateral ocellus contiguous with inner orbit. Upper frons: convex, without frontal shelf. Scrobe shape: frons broadly convex, without distinct scrobe. Frons sculpture: scrobe largely smooth, otherwise granulate or variably punctate. Submedian carina: absent. Orbital carina: absent. Inner orbits: diverging ventrally. IOS/EH: IOS distinctly less than EH; IOS slightly greater than EH. Interantennal process: short, often excavate medially. Central keel: present; absent. Torulus opening: laterally on interantennal process. Lower frons striae: absent. Malar sulcus: present. Compound eye size: of normal proportions, not significantly reduced. Compound eye setation: glabrous; sparsely setose; densely setose. Gena: broad, convex, distinctly produced behind eye. Clypeus shape: narrow, slightly convex medially, lateral corner not produced. Apical margin of clypeus: straight. Anteclypeus: absent. Postclypeus: absent. Labrum: not visible. Mandible shape: moderate. Mandibular teeth: apex with 3, acute, subequal teeth. Arrangement of mandibular teeth: transverse. Number of maxillary palpomeres: 4. Shape of maxillary palpomeres: cylindrical. Number of labial palpomeres: 2. Antenna. 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: A5 only. Shape of male flagellum: filiform. Mesosoma. Mesosoma shape in dorsal view: longer than wide. Mesosoma shape in lateral view: longer than high. Medial portion of transverse pronotal carina: weakly indicated laterally; absent. Posterior apex of pronotum in dorsal view: straight, bifid apically to articulate with tegula. Vertical epomial carina: absent. Dorsal epomial carina (lateral portion of transverse pronotal carina of Vilhelmsen et al. 2010): present. Anterior face of pronotum: oblique, visible dorsally, short. Lateral face of pronotum: weakly concave below position of dorsal epomial carina. Netrion: present. Netrion shape: narrow to moderately wide, open ventrally. Anterior portion of mesoscutum: vertical, flexed ventrally to meet pronotum. Mesoscutum shape: semielliptical, excavate at base of wings. Skaphion: absent. Notauli: present, percurrent; present, abbreviated. Parapsidal lines: absent. Admedial lines: absent. Transscutal articulation: well- developed, narrow. Shape of mesoscutellum: quadrate to trapezoidal. Armature of

16 mesoscutellum: absent. Surface of mesoscutellum: convex throughout. Median longitudinal furrow on mesoscutellum: absent. Shape of axillula: small, dorsal margin sinuate. Metascutellum: clearly differentiated. Metascutellar armature: absent. 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; absent or strongly abbreviated, present only near mid coxa. Mesal course of acetabular carina: projecting as small spur anteriorly, not separating fore coxae. Mesopleural pit: present. Sternaulus: absent. Posterodorsal corner of mesopleuron: rounded anteriorly. Legs. Number of mid tibial spurs: 1. Number of hind tibial spurs: 1. Dorsal surface of hind coxa: smooth; transversely rugose. Hind tibia shape: cylindrical, ecarinate. Trochantellus: indicated by transverse sulcus on femur. Wings. 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: spectral; nebulous, strongly pigmented; absent. Development of R in hind wing: elongate, extending to costal margin. Metasoma. 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: lanceolate. Laterotergites: present, narrow. Laterosternites: present. T1 of female: more or less evenly convex; produced medially into cylindrical or elliptical horn housing ovipositor. Relative size of metasomal segments: T2–T4 largest, subequal in size. Terga with basal crenulae: T2. Sublateral carinae on tergites: absent. Median longitudinal carina on metasomal terga: absent. Shape of female T6: flattened. Shape of posterior margin of male T7: rounded. Anterior margin of S1: not produced anteriorly, concave. Distribution of felt fields: absent. Ovipositor type: Scelio-type (Austin & Field 1997).

Diagnosis. Calliscelio may be distinguished from other genera of the subfamily by the combination of the following characters: eyes glabrous in many species but in some with short hairs or even densely hairy; skaphion never developed; metanotum medially produced into a

17 transverse plate or lamella, neither spinose nor toothed laterally; propodeum usually unarmed, often excavate to contain T1 horn, only in a few species with posterolateral corner acute; T6 in females often elongate, sword-like, depressed dorsoventrally. Calliscelio is most similar to Holoteleia Kieffer and Probaryconus Kieffer in the tribe Calliscelionini and Calotelea in Psilanteridini in body shape and some external characters. The following key is used to separate these genera with the fewest characters possible.

Key to separate Calliscelio, Calotelea, Holoteleia and Probaryconus

1 Skaphion indicated posteriorly by more or less distinct rim ...... Calotelea Westwood - Skaphion never developed, no rim posteriorly ...... 2 2 Genal striae present; epomial carina present ...... Probaryconus Kieffer - Genal striae absent; epomial carina absent ...... 3 3 Metanotum medially notably wider than at sides, expanded into lamella ...... Calliscelio Ashmead - Metanotum narrow, strip-like, medially not produced into lamella ...... Holoteleia Kieffer

New World species of Calliscelio Ashmead

C. absconditum Chen & Johnson, sp. n. C. absum Chen & Johnson, sp. n. C. alcoa Chen & Masner, sp. n. C. amadoi Chen & Johnson, sp. n. C. armila Chen & Masner, sp. n. C. bidens Chen & Masner, sp. n. C.bisulcatus (Kieffer, 1910) C. brachys Chen & Johnson, sp. n. C. brevinotaulus Chen & Johnson, sp. n. C. brevitas Chen & Johnson, sp. n. C. carinigena Chen & Johnson, sp. n. C. crater Chen & Johnson, sp. n.

18 C. crena Chen & Johnson, sp. n. C. eboris Chen & Johnson, sp. n. C. elegans (Perkins) Calotelea tanugatra Narendran C. extenuatus Chen & Johnson, sp. n. C. flavicauda Chen & Johnson, sp. n. C. foveolatus Chen & Johnson, sp. n. C. gatineau Chen & Johnson, sp. n. C. glaber Chen & Masner, sp. n. C. granulatus Chen & Masner, sp. n. C. laticinctus Ashmead, 1893 C. latifrons Chen & Johnson, sp. n. C. levis Chen & Johnson, sp. n. C. longius Chen & Johnson, sp. n. C. magnificus Chen & Masner, sp. n. C. migma Chen & Johnson, sp. n. C. minutia Chen & Johnson, sp. n. C. paraglaber Chen & Johnson, sp. n. C. pararemigio Chen & Masner, sp. n. C. prolixus Chen & Johnson, sp. n. C. punctatifrons Chen & Johnson, sp. n. C. remigio Chen & Masner, sp. n. C. rubriclavus (Ashmead, 1887), comb. n. Anteris nigriceps Ashmead, 1893, syn. n. Caloteleia Marlattii Ashmead, 1893, syn. n. Caloteleia grenadensis Ashmead, 1896, syn. n. Macroteleia ruskini Girault, 1920, syn. n. C. ruga Chen & Johnson, sp. n. C. rugicoxa Chen & Masner, sp. n. C. sfina Chen & Johnson, sp. n. C. storea Chen & Johnson, sp. n.

19 C. suni Chen & Johnson, sp. n. C. telum Chen & Johnson, sp. n. C. torqueo Chen & Johnson, sp. n. C. virga Chen & Johnson, sp. n.

Key to Females of Calliscelio of the New World

1 Occipital carina complete medially (Figs 1.3D, 1.6D, 1.7D, 1.8D, 1.9D, 1.12D, 1.15D, 1.17D, 1.18D, 1.19D, 1.20D, 1.22D, 1.23D, 1.24D, 1.25D, 1.26D, 1.27D, 1.28D, 1.30D, 1.32D, 1.33D, 1.36D, 1.39D, 1.41D, 1.43D, 1.45D) ...... 2 - Occipital carina interrupted medially (Figs 1.4D, 1.5D, 1.10D, 1.11D, 1.13D, 1.14D, 1.16D, 1.21D, 1.29D, 1.31D, 1.34D, 1.38D, 1.40D, 1.42D, 1.44D) ...... 28 2 Eye bare (Figs 1.6E, 1.12E, 1.15E, 1.18E, 1.19E, 1.20E, 1.23E, 1.25E, 1.26E, 1.27E, 1.28E, 1.30E, 1.32E, 1.36E) ...... 3 - Eye setose (Figs 1.7E, 1.9E, 1.8E, 1.17E, 1.22D, 1.24E, 1.33E, 1.39E, 1.41E, 1.43E, 1.45E) ...... 17 3 A4 distinctly shorter than A3 (Figs 1.3C, 1.6C, 1.12C, 1.15C, 1.19C, 1.20A, 1.30E) ...... 4 - A4 approximately as long as or distinctly longer than A3 (Figs 1.1J, 1.23E, 1.25C, 1.27C, 1.28A, 1.32A) ...... 11 4 Horn on T1 weakly developed, smooth (Figs 1.19F, 1.30F) ...... 5 - Horn on T1 large and distinct, variably sculptured (Figs 1.3F, 1.6F, 1.20F, 1.36D) ...... 6 5 Metascutellum without a longitudinal median carina (Fig. 1.30D); foveolae of scutoscutellar sulcus between notauli smaller than those along margin of axilla (Fig. 1.30D); posterior vertex smooth throughout (Fig. 1.30D) ...... Calliscelio paraglaber Chen & Johnson, sp. n. - Metascutellum with a longitudinal median carina (Fig. 1.19D); foveolae of scutoscutellar sulcus between notauli as large as those along margin of axilla (Fig. 1.19D); posterior vertex smooth to coriaceous (Fig. 1.19D) ...... Calliscelio foveolatus Chen & Johnson, sp. n. 6 T6 strongly elongate, at least 2.0x longer than wide (Figs 1.10F, 1.15F) ...... 7 - T6 short, at most 1.5x longer than wide (Figs 1.3F, 1.6F, 1.20F, 1.37E) ...... 8

20 7 Setae on frons short (Fig. 1.12E); posterior vertex granulate to rugulose or densely punctate (Fig. 1.12D); mesoscutum largely coriaceous with dense and fine punctures at posterior extreme (Fig. 1.12D) ...... Calliscelio brevitas Chen & Johnson, sp. n. - Setae on frons long (Fig. 1.15E); posterior vertex largely smooth with sparse fine punctures (Fig. 1.15D); mesoscutum with anterior margin rugulose, remainder smooth (Fig. 1.15D) ...... Calliscelio crena Chen & Johnson, sp. n. 8 R1 as long as r-rs (Figs 1.2A, 1.35E); IOS slightly less than or greater than EH (Figs 1.20E, 1.36E) ...... 9 - R1 as long as 2.0x length of r-rs (Figs 1.3F, 1.6F); IOS distinctly less than EH (Figs 1.3E, 1.6E) ...... 10 9 Mesepisternum below mesopleural depression densely punctate (Fig. 1.35D); horn on T1 large and distinct, granulate or rugose dorsally (Fig. 1.36D) ...... Calliscelio rubriclavus (Ashmead) - Mesepisternum below mesopleural depression smooth (Fig. 1.20B); horn on T1 present as a small bulge, dorsally granulate medially, with V-shaped keels on edge (Fig. 1.20F) ...... Calliscelio gatineau Chen & Johnson, sp. n. 10 Posterior vertex largely smooth with sparse fine punctures (Fig. 1.6D); Rs+M dark, nebulous (Fig. 1.6F); T1 horn transversely striate (Fig. 1.6F) ...... Calliscelio amadoi Chen & Johnson, sp. n. - Posterior vertex granulate to rugulose (Fig. 1.3D); Rs+M spectral (Fig. 1.3F); T1 horn with V-shaped striae (Fig. 1.3F) ...... Calliscelio absconditum Chen & Johnson, sp. n. 11 T6 short, at most 1.5x longer than wide (Figs 1.18F, 1.25F, 1.28F, 1.32F) ...... 12 - T6 strongly elongate, at least 2.0x longer than wide (Figs 1.26F, 1.27F) ...... 15 12 Horn on T1 absent or weakly developed (Figs 1.25F, 1.32F); dorsal propodeum not excavate medially, lateral propodeal carinae meeting anteromedially (Figs 1.25D, 1.32D) ...... 13 - Horn on T1 present as a small bulge (Figs 1.18F, 1.28F); dorsal propodeum shallowly excavate medially, with lateral propodeal carinae widely separated (Fig. 1.18D, 1.28D) ...... 14 13 A4 distinctly longer than A3 (Fig. 1.32C); upper frons densely setose (Fig. 1.32E); T1 longitudinally striate medially (Fig. 1.32F) ...... Calliscelio prolixus Chen & Johnson, sp. n.

21 - A4 as long as A3 (Fig. 1.25C); upper frons sparsely setose (Fig. 1.25E); T1 smooth medially (Fig. 1.25F) ...... Calliscelio levis Chen & Johnson, sp. n. 14 Scutoscutellar sulcus strongly foveolate medially (Fig. 1.18D); upper frons densely setose (Fig. 1.18E); T3 smooth throughout (Fig. 1.18F)...... Calliscelio flavicauda Chen & Johnson, sp. n. - Scutoscutellar sulcus weakly foveolate medially (Fig. 1.28D); upper frons sparsely setose (Fig. 1.28E); T3 with longitudinal submedian striae (Fig. 1.28F) ...... Calliscelio migma Chen & Johnson, sp. n. 15 Horn on T1 smooth (Fig. 1.27F); foveolae of scutoscutellar sulcus between notauli smaller than those along margin of axilla (Fig. 1.27D) ...... Calliscelio magnificus Chen & Masner, sp. n. - Horn on T1 at least partly transversely striate (Figs 1.23D, 1.26F); foveolae of scutoscutellar sulcus between notauli as large as those along margin of axilla (Fig. 1.23D, 1.26D) ...... 16 16 A4 distinctly longer than A3 (Fig. 1.1J); A5 longer than A3 (Fig. 1.1J) ...... Calliscelio longius Chen & Johnson, sp. n. - A4 slightly longer than A3 (Fig. 1.23E); A5 shorter than A3 (Fig. 1.23E) ...... Calliscelio laticinctus Ashmead 17 Median keels on propodeum present (Figs 1.8D, 1.39D)...... 18 - Median keels on propodeum absent (Figs 1.7D, 1.9D, 1.17D, 1.22D, 1.24D, 1.33D, 1.41D, 1.43D, 1.45D) ...... 19 18 Horn on T1 absent, anterior margin of T1 longitudinally striate (Fig. 1.8F); hind coxae smooth (Fig. 1.8B); T6 short, slightly longer than wide (Fig. 1.8F) ...... Calliscelio bidens Chen & Masner, sp. n. - Horn on T1 weakly developed, rugose dorsally (Fig. 1.39F); hind coxae rugose (Fig. 1.39B); T6 distinctly elongate, 2.0x longer than wide (Fig. 1.39F) ...... Calliscelio rugicoxa Chen & Masner, sp. n. 19 T6 strongly elongate, at least 2.0x longer than wide (Figs 1.22F, 1.43F) ...... 20 - T6 short, at most 1.5x longer than wide (Figs 1.7F, 1.9F, 1.17F, 1.24F, 1.33F, 1.41F, 1.45F) ...... 22 20 Fore wing strikingly banded, with dark bands basally, medially and apically, separated by light bands; R1 only slightly longer than r-rs ...... Calliscelio elegans (Perkins)

22 - Fore wing hyaline; R1 approximately as long as 2.0x length of r-rs ...... 21 21 Setae of upper frons short, dense (Fig. 1.22E); netrion rugose (Fig. 1.22B); horn on T1 longitudinally striate (Fig. 1.22F) ...... Calliscelio granulatus Chen & Johnson, sp. n. - Setae of upper frons long, sparse (Fig. 1.43E); netrion smooth (Fig. 1.43B); horn on T1 rugulose (Fig. 1.43F) ...... Calliscelio telum Chen & Johnson, sp. n. 22 Horn on T1 absent (Fig. 1.17F); dorsal propodeum not excavate medially (Fig. 1.17D) ...... Calliscelio extenuates Chen & Johnson, sp. n. - Horn on T1 present (Figs 1.7F, 1.9F, 1.24F, 1.33F, 1.41F, 1.45F); dorsal propodeum excavate medially (Figs 1.7D, 1.9D, 11.7F, 1.33D, 1.41D, 1.45D) ...... 23 23 Horn on T1 variably sculptured (Figs 1.7F, 1.24F, 1.41F) ...... 24 - Horn on T1 smooth (Figs 1.9F, 1.33F, 1.45F) ...... 26 24 Notaulus abbreviated, at most reaching middle of mesoscutum (Fig. 1.24D); T3 with submedian longitudinal striae (Fig. 1.24F)...... Calliscelio latifrons Chen & Johnson, sp. n. - Notaulus percurrent (Figs 1.7D, 1.41D); T3 smooth throughout (Figs 1.7F, 1.41F) ...... 25 25 Horn on T1 rugulose (Fig. 1.41F); legs orange throughout (Fig. 1.41A)...... Calliscelio storea Chen & Johnson, sp. n. - Horn on T1 smooth to rugulose medially, with V-shaped keels laterally (Fig. 1.7F); legs with coxae and femora white, otherwise pale yellow throughout (Fig. 1.7A) ...... Calliscelio armila Chen & Masner, sp. n. 26 Frons below median ocellus densely punctate (Fig. 1.33E); posterior vertex densely punctate (Fig. 1.33D); R distinctly longer than r-rs (Fig. 1.33F) ...... Calliscelio punctatifrons Chen & Johnson, sp. n. - Frons below median ocellus largely smooth with sparse fine punctures (Figs 1.9E, 1.45E); posterior vertex granulate (Figs 1.9D, 1.45D); R approximately as long as r-rs (Fig. 1.45A) ...... 27 27 Metascutellum approximately 4.0x wider than long, smooth (Fig. 1.45D); mesopleural carina absent (Fig. 1.45B); mesepisternum below mesopleural depression smooth throughout (Fig. 1.45B) ...... Calliscelio virga Chen & Johnson, sp. n. - Metascutellum approximately 2.0x wider than long, rugose anteriorly, smooth posteriorly (Fig. 1.9D); mesopleural carina present (Fig. 1.9B); mesepisternum below mesopleural

23 depression with a row of foveae along mesopleural carina ...... Calliscelio bisulcatus (Kieffer) 28 Horn on T1 absent or at most weakly developed (Figs 1.4F, 1.10F, 1.16F, 1.29F, 1.43F)29 - Horn on T1 present (Figs 1.5F, 1.11F, 1.14F, 1.21F, 1.31F, 1.34F, 1.40F, 1.44F) ...... 34 29 Notaulus abbreviated, at most reaching middle of mesoscutum (Figs 25, 1.10D) ...... 30 - Notaulus percurrent (Figs 1.16D, 1.29D, 1.38D, 1.43D) ...... 31 30 A4 distinctly shorter than A3 (Fig. 1.4A); postgena behind outer orbit coriaceous (Fig. 1.4B) ...... Calliscelio absum Chen & Johnson, sp. n. - A4 as long as A3 (Fig. 1.10C); postgena behind outer orbit smooth (Fig. 1.10B) ...... Calliscelio brachys Chen & Johnson, sp. n. 31 Eye hairy (Fig. 1.38D); A6 distinctly longer than wide ...... Calliscelio ruga Chen & Johnson, sp. n. - Eye bare (Figs 1.16E, 1.29E, 1.43E); A6 subquadrate or distinctly transverse ...... 32 32 Fore wing hyaline with an infuscate band in the middle (Fig. 1.29C); A4 distinctly shorter than A3; postgena behind outer orbit granulate (Fig. 1.29B) ...... Calliscelio minutia Chen & Johnson, sp. n. - Fore wing entirely hyaline (Figs 1.16F, 1.43C); A4 as long as A3; postgena behind outer orbit smooth (Figs 1.16B, 1.43B) ...... 33 33 Central keel of frons absent (Fig. 1.43E); frons below median ocellus smooth (Fig. 1.43E); Rs+M nebulose, weakly pigmented (Fig. 1.43F) .... Calliscelio suni Chen & Johnson, sp. n. - Central keel of frons present (Fig. 1.16E); frons below median ocellus coriaceous (Fig. 1.16E); Rs+M spectral (Fig. 1.16F) ...... Calliscelio eboris Chen & Johnson, sp. n. 34 Frons and posterior vertex smooth (Figs 1.21D, 1.21E, 1.31D, 1.31E, 1.34D, 1.34E, 1.44D, 1.44E) ...... 35 - Frons and posterior vertex variably sculptured (Figs 1.5D, 1.5E, 1.11D, 1.11E, 1.13D, 1.13E, 1.14D, 1.14E, 1.40D, 1.40E) ...... 38 35 T6 strongly elongate, approximately 3.0x longer than wide (Figs 1.31F, 1.34F); Metascutellum finely granulate (Figs 1.31D, 1.34D) ...... 36 - T6 short, slightly longer than wide (Figs 1.21F, 1.44F); Metascutellum finely smooth (Figs 1.21D, 1.44D) ...... 37

24 36 Head strongly transverse in dorsal view (Fig. 1.34D); T1 horn transversely striate (Fig. 1.34F)...... Calliscelio remigio Chen & Masner, sp. n. - Head subglobose in dorsal view (Fig. 1.31D); T1 horn densely and concentrically striate anteriorly, smooth posteriorly (Fig. 1.31F) ...... Calliscelio pararemigio Chen & Masner, sp. n. 37 A4 distinctly longer than A3 (Fig. 1.44C); horn on T1 smooth (Fig. 1.44F) ...... Calliscelio torque Chen & Johnson, sp. n. - A4 distinctly shorter than A3 (Fig. 1.21C); horn on T1 concentrically striate (Fig. 1.21F) ...... Calliscelio glaber Chen & Masner, sp. n. 38 Hyperoccipital carina present (Figs 1.13D, 1.14D) ...... 39 - Hyperoccipital carina absent (Figs 1.5D, 1.11D, 1.40D) ...... 40 39 Postgena behind outer orbit with large foveae (Fig. 1.14B); horn on T1 large and distinct, rugose medially, with V-shaped keels laterally (Fig. 1.14F) ...... Calliscelio crater Chen & Johnson, sp. n. - Postgena with a carina along outer orbit (Fig. 1.13B); horn on T1 weakly indicated, smooth to somewhat transversely striate (Fig. 1.11F) ...... Calliscelio carinigena Chen & Johnson, sp. n. 40 Mesoscutum densely punctate (Fig. 1.5D); horn on T1 densely and transversely striate (Fig. 1.5F); T6 strongly elongate, tapering apically (Fig. 1.5F) ...... Calliscelio alcoa Chen & Masner, sp. n. - Mesoscutum granulate (Figs 1.11D, 1.40D); horn on T1 with V-shaped striae (Figs 1.11F, 1.40F); T6 short, subtriangular (Figs 1.11F, 1.40F) ...... 41 41 Notaulus abbreviated, at most reaching anteriorly to middle of mesoscutum (Fig. 1.11D); ventral metapleural area largely smooth (Fig. 1.11B); S3 largely smooth with sparse fine punctures...... Calliscelio brevinotaulus Chen & Johnson, sp. n. - Notaulus percurrent (Fig. 1.40D); ventral metapleural area rugose throughout (Fig. 1.40B); S3 densely punctate medially, longitudinally striate laterally ...... Calliscelio sfina Chen & Johnson, sp. n.

Key to Males (unknown for C. amadoi, C. bidens, C. brevitas, C. foveolatus, C. gatineau, C. levis, C. prolixus, C. rugicoxa, C. ruga and C. storea)

25

1 Occipital carina complete medially (Figs 1.3D, 1.7D, 1.9D, 1.15D, 1.17D, 1.18D, 1.22D, 1.23D, 1.24D, 1.26D, 1.27D, 1.28D, 1.30D, 1.33D, 1.36D, 1.43D, 1.45D) ...... 2 - Occipital carina interrupted medially (Figs 1.4D, 1.5D, 1.10D, 1.11D, 1.13D, 1.14D, 1.16D, 1.21D, 1.29D, 1.31D, 1.34D, 1.40D, 1.43D, 1.44D) ...... 19 2 Eye bare (Figs 1.3E, 1.15E, 1.18E, 1.23E, 1.26E, 1.27E, 1.28E, 1.30E) ...... 3 - Eye setose (Figs 1.7E, 1.17D, 1.22D, 1.33E, 1.43E, 1.45E) ...... 11 3 Rs+M spectral (Figs 1.3F, 1.35E); R distinctly shorter than r-rs ...... 4 - Rs+M nebulose, pigmented (Figs 1.2C, 1.15C, 1.26F, 1.27F, 1.28F, 1.30F); R as long as or longer than r-rs ...... 5 4 Mesepisternum below mesopleural depression densely punctate (Fig. 1.35D); IOS slightly less than EH (Fig. 1.36E); R1 as long as r-rs ...... Calliscelio rubriclavus (Ashmead) - Mesepisternum below mesopleural depression smooth (Fig. 1.3B); IOS distinctly less than EH (Fig. 1.3E); R1 approximately as long as 2.0x length of r-rs ...... Calliscelio absconditum Chen & Johnson, sp. n. 5 Upper frons densely setose (Figs 1.18E, 1.23E, 1.26E, 1.27E)...... 6 - Upper frons sparsely setose (Figs 1.15E, 1.28E, 1.30E) ...... 9 6 Metascutellum smooth (Fig. 1.27D) ...... Calliscelio magnificus Chen & Masner, sp. n. - Metascutellum rugose (Figs 1.18D, 1.23D, 1.26D) ...... 7 7 A11 approximately 4.5x longer than wide .. Calliscelio longuis Chen & Johnson, sp. n. - A11 approximately 3.0x longer than wide ...... 8 8 T3 smooth throughout (Fig. 1.18F); hind femora brown ...... Calliscelio flavicauda Chen & Johnson, sp. n. - T3 with longitudinal submedian striae; hind femora yellow ...... Calliscelio laticinctus Ashmead 9 A11 approximately 2.0x longer than wide; length of T5 tyloid greater than 0.5x length of A5 ...... Calliscelio crena Chen & Johnson, sp. n. - A11 approximately 4.0x longer than wide; length of T5 tyloid approximately 0.3x length of A5 ...... 10 10 Mesoscutum smooth throughout (Fig. 1.30D); T3 smooth throughout (Fig. 1.30F) ...... Calliscelio paraglaber Chen & Johnson, sp. n.

26 - Mesoscutum coriaceous or smooth with sparse punctures (Fig. 1.28D); T3 with longitudinal submedian striae (Fig. 1.28F) ...... Calliscelio migma Chen & Johnson, sp. n. 11 R1 approximately as long as r-rs ...... 12 - R1 at least 2.0x length of r-rs ...... 14 12 Fore wing strikingly banded, with dark bands basally, medially and apically, separated by light bands ...... Calliscelio elegans (Perkins) - Fore wing hyaline (Figs 1.9C, 1.45A) ...... 13 13 Metascutellum rugose (Fig. 1.9D); mesepisternum below mesopleural depression largely smooth with a row of foveae along mesopleural carina ...... Calliscelio bisulcatus (Kieffer) - Metascutellum smooth (Fig. 1.45D); mesepisternum below mesopleural depression smooth throughout (Fig. 1.45B) ...... Calliscelio virga Chen & Johnson, sp. n. 14 IOS slightly greater than EH (Fig. 1.24E); antennal flagellomeres moniliform, A11 as long as wide (Fig. 1.1E)...... Calliscelio latifrons Chen & Johnson, sp. n. - IOS less than EH (Figs 1.7E, 1.17E, 1.22E, 1.33E, 1.43E); antennal flagellomeres filiform, A11 at least 2.5x longer than wide (Fig. 1.1B) ...... 15 15 Posterior vertex densely punctate (Fig. 1.33D); mesostutellum smooth with sparse fine punctures (Fig. 1.33D) ...... Calliscelio punctatifrons Chen & Johnson, sp. n. - Posterior vertex granulate (Figs 1.7D, 1.17D, 1.22D, 1.43D); mesoscutellum granulate (Figs 1.7D, 1.17D, 1.22D, 1.43D) ...... 16 16 R1 greater than 3.0x length of r-rs...... Calliscelio extenuates Chen & Johnson, sp. n. - R1 approximately as long as 2.0x length of r-rs ...... 17 17 Frons below median ocellus largely smooth with sparse fine punctures (Fig. 1.7E); IOS slightly less than EH (Fig. 1.7E) ...... Calliscelio armila Chen & Masner, sp. n. - Frons below median ocellus granulate (Figs 1.22E, 1.43E); IOS distinctly less than EH (Figs 1.22E, 1.43E) ...... 18 18 Rs+M nebulose, strongly pigmented; dorsal propodeum with one or two longitudinal keels lateral to median keels ...... Calliscelio telum Chen & Johnson, sp. n. - Rs+M spectral; dorsal propodeum rugose throughout, only with medial keels ...... Calliscelio granulatus Chen & Masner, sp. n. 19 Notaulus abbreviated, at most reach anteriorly to middle of mesoscutum (Figs 1.4D, 1.10D) ...... 20

27 - Notaulus percurrent (Figs 1.5D, 1.13D, 1.14D, 1.16D, 1.21D, 1.29D, 1.31D, 1.34D, 1.40D, 1.43D, 1.44D) ...... 22 20 Netrion rugose (Fig. 1.11B); mesopleural carina absent (Fig. 1.11B) ...... Calliscelio brevinotaulus Chen & Johnson, sp. n. - Netrion smooth (Figs 23, 1.10B); mesopleural carina present (Figs 23, 1.10B) ...... 21 21 Postgena behind outer orbit smooth (Fig. 1.10B); length of A5 tyloid approximately 0.3x length of A5; dorsal propodeum with one or two longitudinal keels lateral to median keel ...... Calliscelio brachys Chen & Johnson, sp. n. - Postgena behind outer orbit coriaceous (Fig. 1.4B); length of A5 tyloid longer than 0.5x length of A5; lateral propodeal area rugose throughout ...... Calliscelio absum Chen & Johnson, sp. n. 22 Frons below median ocellus smooth (Figs 1.21E, 1.31E, 1.34E, 1.43E, 1.44E) ...... 23 - Frons below median ocellus sculptured (Figs 1.5E, 1.13E, 1.14E, 1.16E, 1.29E, 1.40E)27 23 Rs+M nebulose, pigmented (Figs 1.43C, 1.44C) ...... 24 - Rs+M spectral (Figs 1.21C, 1.31C, 1.34C) ...... 25 24 Metascutellum rugose (Fig. 1.43D); R distinctly shorter than r-rs; T3 smooth throughout (Fig. 1.43F) ...... Calliscelio suni Chen & Johnson, sp. n. - Metascutellum smooth (Fig. 1.44D); R approximately as long as r-rs; T3 with longitudinal submedian striae (Fig. 1.44F) ...... Calliscelio torqueo Chen & Johnson, sp. n. 25 Mesopleural carina absent (Fig. 1.21B); mesoscutellum smooth throughout (Fig. 1.21D) ...... Calliscelio glaber Chen & Masner, sp. n. - Mesopleural carina present (Figs 1.31B, 1.34B); mesoscutellum with sparse fine punctures (Fig. 1.31D, 1.34D) ...... 26 26 Head strongly transverse in dorsal view (Fig. 1.34D); S3 densely punctate or punctate rugose (Fig. 1.34F) ...... Calliscelio remigio Chen & Masner, sp. n. - Head subglobose in dorsal view (Fig. 1.31D); S3 largely smooth with sparse fine punctures (Fig. 1.31F) ...... Calliscelio pararemigio Chen & Masner, sp. n. 27 Mesoscutum densely punctate (Fig. 1.5D); R approximately as long as r-rs; ...... Calliscelio alcoa Chen & Masner, sp. n. - Mesoscutum coriaceous or granulate (Figs 1.13D, 1.14D, 1.16D, 1.29D, 1.40D); R distinctly shorter than r-rs; ...... 28

28 28 Hyperoccipital carina present (Figs 1.13D, 1.14D, 1.16D) ...... 29 - Hyperoccipital carina absent (Figs 1.29D, 1.40D) ...... 31 29 Postgena behind outer orbit with large foveae (Fig. 1.14B) ...... Calliscelio crater Chen & Johnson, sp. n. - Postgena behind outer orbit without foveae (Figs 1.13B, 1.16B) ...... 30 30 Posterior vertex transversely striate (Fig. 1.16D); postgena behind outer orbit smooth (Fig. 1.16B) ...... Calliscelio eboris Chen & Johnson, sp. n. - Posterior vertex granulate to rugulose (Fig. 1.13D); postgena behind outer orbit with a carina along outer orbit (Fig. 1.13B) ...... Calliscelio carinigena Chen & Johnson, sp. n. 31 S3 smooth (Fig. 1.29F); fore wing hyaline with infuscate band in the middle (Fig. 1.29C); ventral metapleural area smooth dorsally, densely punctate ventrally (Fig. 1.29B) ...... Calliscelio minutia Chen & Johnson, sp. n. - S3 densely punctate medially, longitudinally striate laterally (Fig. 1.40F); fore wing hyaline throughout (Fig. 1.40C); ventral metapleural area rugose (Fig. 1.40B) ...... Calliscelio sfina Chen & Johnson, sp. n.

Species descriptions

Calliscelio absconditumChen & Johnson, new species http://zoobank.org/7A1C9B4D-B25D-403F-9DE3-B2278A4BFED8 http://bioguid.osu.edu/xbiod_concepts/384359 Figures 1.3A–1.3F.

Description. Body length of female: 2.04–2.54 mm (n=20). Body length of male: 1.70–2.36 mm (n=20). Color of head: variably brown to black. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth to coriaceous. Sculpture of frons below median ocellus: coriaceous. Sculpture of posterior vertex: granulate to rugulose. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: largely smooth with small granulate area. Ocular setae: absent. A4

29 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Form of male antennal flagellomeres: filiform, A11 approximately 2.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: orange throughout; variably orange to pale brown. Color of mesosoma in male: variably orange to pale brown; dark brown throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 3.0x wider than long. Sculpture of metascutellum in female: rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth with an oblique carina. Color of legs: pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: orange throughout; variably orange to pale brown. Color of metasoma in male: brown throughout. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: smooth to rugulose medially, with V-shaped keels laterally; with V-shaped striae. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth; smooth medially, coriaceous laterally. Shape of T6 in female: short, wider than long. Sculpture of S3: smooth to coriaceous. Diagnosis. This species is similar to C. brevinotaulus, C. carinigena, C. crater and C. sfina in color pattern, size, and habitus. It may be distinguished by the complete occipital carina and granulate to rugulose posterior vertex (Fig. 1.3D).

30 Etymology. The epithet is used as a noun in apposition derived from the Latin word for hidden, in reference to the obscurity of the occipital carina from dorsal view. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384359] Material Examined. Holotype, female: PARAGUAY: Canindeyú Dept., Jejuí-mí, Bosque Mbaracayú Natural Reserve, 10.I.1997, B. Garcete, OSUC 323924 (deposited in MNHNPY). Paratypes: (59 females, 104 males) BOLIVIA: 7 females, OSUC 534032, 534038, 534040, 534054–534055, 534058, 534186 (CNCI). BRAZIL: 11 females, 28 males, OSUC 534520 (CNCI); OSUC 10439, 10444, 10601, 10610, 10653, 110189, 131761, 131787, 131827, 134436, 134786, 134875, 510887, 826 (MZSP); OSUC 10040, 10502, 10557, 10633, 10718, 10949, 110085, 110139, 110159, 110215, 11065, 112, 12321, 130, 131695, 131754, 131816, 131840, 131892, 133022, 133090, 134370, 134681, 134791 (OSUC). PARAGUAY: 41 females, 76 males, OSUC 534107-534115, 534559 (CNCI); OSUC 534683, 534695-534697, 534699- 534700, 534702, 534704, 534706-534707, 534713-534714, 534726-534728, 534731-534749, 570521-570525, 570527-570534 (MNHNPY); OSUC 150602-150603, 150606, 150610-150611, 165099, 176064, 276773-276777, 276796, 278657-278658, 278661-278662, 278664-278665, 278668-278670, 278673, 278676, 278679, 322990, 323001-323003, 323027-323029, 323032, 323035, 323920-323923, 323925-323927, 412079-412082, 412085, 534725, 570526, 577174, 577176, 577178-577180, 577188-577189, 577340, 577343, 577346, 577349-577350 (OSUC).

Calliscelio absumChen & Johnson, new species http://zoobank.org/2877EC6A-9602-499F-9075-0CCC91166CD9 http://bioguid.osu.edu/xbiod_concepts/384780 Figures 1.4A–1.4F.

Description. Body length of female: 1.49–2.58 mm (n=20). Body length of male: 1.49–1.60 mm (n=20). Color of head: black throughout; dark brown; orange throughout. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth to granulate. Sculpture of frons below median ocellus: smooth; coriaceous. Sculpture of posterior vertex: smooth; coriaceous. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture

31 of postgena behind outer orbit: coriaceous. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Form of male antennal flagellomeres: filiform, A11 approximately 2.5x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: orange throughout; black throughout; orange to pale brown. Color of mesosoma in male: orange throughout; dark brown throughout; black throughout. Sculpture of dorsal pronotal area: smooth. Sculpture of lateral pronotal area: smooth throughout; smooth anteriorly, granulate posteriorly. Sculpture of netrion: smooth. Notaulus: abbreviated, at most reaching middle of mesoscutum. Sculpture of mesoscutum: coriaceous. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: coriaceous; anterior half granulate, posterior half smooth. Shape of metascutellum: posterior margin rounded, approximately 3.0x wider than long. Sculpture of metascutellum in female: with short longitudinal carinae. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth with an oblique carina. Color of legs: pale yellow throughout; white throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: greater than 3.0x length of r- rs. Color of metasoma in female: dark brown; orange throughout; yellow throughout. Color of metasoma in male: orange throughout; brown throughout. Horn on T1 in female: absent; weakly developed. Sculpture of T1 horn dorsally: smooth. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially; reaching posterior margin of T2. Sculpture of T3: smooth; smooth medially, longitudinally striate laterally. Shape of T6 in female: short, wider than long. Sculpture of S3: smooth.

32 Diagnosis. This species is most similar to C. brachys in size and abbreviated notaulus. It may be distinguished from C. brachys in having A4 distinctly shorter than A3, A6 distinctly transverse in the female. Etymology. The specific epithet is to be treated as a noun in apposition, derived from the Latin for "be away," and refers to the abbreviated notaulus. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384780] Material Examined. Holotype, female: COLOMBIA: Magdalena Dept., Zaino, M.567, 50m, 11°20'N 74°02'W, Tayrona Natural National Park, 28.VII–14.VIII.2000, Malaise trap, R. Henriquez, OSUC 190716 (deposited in IAVH). Paratypes: (146 females, 106 males) BELIZE: 42 females, 27 males, OSUC 534029 (CNCI); OSUC 185847, 185849–185851, 185854, 237727–237729, 246215–246217, 247494, 342738–342741, 397563, 398893, 47953, 47963, 47971, 48049–48052, 48054, 64034, 64038, 64041, 64043, 64073, 64077, 64090, 64096–64097, 64102, 64109, 64111, 64116, 91676, 91685, 91687, 91689, 91693, 91695–91696, 91699–91700, 91702, 93079, 93526, 93590–93593, 93714, 93735, 93743, 93745, 94041–94042, 94046–94047, 94083–94084, 94087–94089 (OSUC). COLOMBIA: 75 females, 59 males, OSUC 557540- 557545, 557558, 557560, 557574-557575, 557586, 557629-557632 (CNCI); OSUC 188886, 188888, 189393-189394, 189397, 190667-190669, 190675-190676, 190679, 192201, 192203, 192206-192207, 192210, 192212-192213, 192217-192223, 192382-192384, 192391-192392, 192395-192396, 192398-192399, 194186-194188, 194190, 194192, 194195 (IAVH); OSUC 170498-170502, 189905, 189907-189909, 190096, 190099, 190101, 190711-190712, 190714, 190717-190718, 190993, 191058-191060, 191063, 191144, 191240, 191244, 191252, 191264- 191266, 191269-191271, 191277, 191279-191280, 191282, 191740, 192208, 192224, 193702- 193703, 193705, 193771, 193773-193776, 193808, 194191, 194197, 194199, 259569-259571, 259578, 259580-259581, 259584-259587, 259590-259591, 279453, 279455-279459, 364087- 364089, 369978, 370059, 370061-370062, 370065-370067 (OSUC).COSTA RICA: 1 female, OSUC 532552 (CNCI). HONDURAS: 12 females, OSUC 399367, 399373, 399383, 399388– 399390, 399406, 399408, 410435, 410440, 410443, 410447 (MZLU). MEXICO: 7 females, 13 males, OSUC 534463, 534465–534470, 534479–534482, 534487 (CNCI); OSUC 55922, 55928, 55937, 55949, 576983–576986 (OSUC). TRINIDAD AND TOBAGO: 2 females, OSUC 546012, 546025 (CNCI). VENEZUELA: 7 females, 7 males, OSUC 545849–545850, 545854, 545856–545857, 545870, 545900–545901, 545975–545978, 557666, 557677 (CNCI).

33

Calliscelio alcoa Chen & Masner, new species http://zoobank.org/A4769E20-E5A0-42CE-8768-D3C39939FC9D http://bioguid.osu.edu/xbiod_concepts/362051 Figures 1.1G–1.1I, 1.5A–1.5F.

Description. Body length of female: 2.86–3.10mm (n=10). Body length of male: 2.42–2.54mm (n=16). Color of head: orange throughout. Color of antennal clava (A7–A12): A7 dark orange, remainder dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with dense, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate to finely punctate. Sculpture of frons below median ocellus: largely smooth with sparse fine punctures. Sculpture of posterior vertex: densely punctate. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: distinctly longer than wide. Form of male antennal flagellomeres: filiform, A11 approximately 3.0x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: orange throughout. Color of mesosoma in male: orange throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth anteriorly, punctate rugulose posteriorly. Sculpture of netrion: rugulose. Notaulus: percurrent. Sculpture of mesoscutum: densely punctate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin somewhat rounded, approximately 4.0x wider than long. Sculpture of metascutellum in female: rugose anteriorly, smooth posteriorly. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral to median keel. Median keels on propodeum in female: absent. Mesopleural carina: present.

34 Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, weakly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: yellow to dark brown. Color of metasoma in male: T3 yellow, otherwise brown to dark brown. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: densely and transversely striate. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth medially, longitudinally striate laterally. Shape of T6 in female: distinctly elongate, approximately 3.5x longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. The medially interrupted occipital carina, densely punctate mesoscutum and densely and transversely striate T1 horn in the female distinguish this species of Calliscelio from all others. Etymology. The specific epithet refers to the locality on the label of the holotype and should be treated as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362051] Material Examined. Holotype, female: DOMINICAN REPUBLIC: Pedernales Prov., 25km of Alcoa Road, dry montane forest, Baoruco (Bahoruco) Mountains, 700m, 18.I.1989, L. Masner, OSUC 458212 (deposited in CNCI). Paratypes: DOMINICAN REPUBLIC: 9 females, 16 males, CMNH-486,529 (CMNH); OSUC 458213–458232, 458251, 458327–458328, 534379 (CNCI).

Calliscelio amadoiChen & Johnson, new species http://zoobank.org/6571E5B9-E640-47B3-AD2C-D654DC26829F http://bioguid.osu.edu/xbiod_concepts/384810 Figures 1.6A–1.6F.

35 Description. Body length of female: 1.35–2.05 mm (n=20). Color of head: brown throughout; yellow throughout; yellow, becoming darker at vertex. Color of antennal clava (A7–A12): A7 orange, remainder dark brown to black. Shape of head: subglobose. Central keel of frons: present. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth with sparse punctures. Sculpture of frons below median ocellus: largely smooth with sparse fine punctures. Sculpture of posterior vertex: largely smooth with sparse fine punctures. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, slightly longer than wide. Shape of female A6: as long as wide. Color of mesosoma in female: orange throughout; yellow throughout; dark brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth anteriorly, granulate posteriorly. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: coriaceous. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: smooth with a longitudinal, median carina. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: orange to pale brown; orange throughout; yellow throughout. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: rugulose; transversely striate. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Development of longitudinal striae on T2 in female: present on anterior margin of T2 medially, reaching posterior margin of T2 laterally. Sculpture of T3: smooth with

36 longitudinal submedian striae. Shape of T6 in female: short, approximately 1.5x longer than wide. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. levis and C. migma. It may be separated from C. levis by the well-developed T1 horn in the female, from C. migma by having A5 in female slightly longer than wide, and A6 quadrate, whereas both antennomeres are distinctly longer than wide in C. migma. Etymology. The epithet is used as a genitive noun derived from the name of the collector of the holotype. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384810] Material Examined. Holotype, female: COLOMBIA: Amazonas Dept., Amacayacu Natural N.P., M.840, 150m, 03°46'S 70°18'W, San Martín, 16.IX–24.IX.2000, Malaise trap, B. Amado, OSUC 193589 (deposited in IAVH). Paratypes: (50 females) BRAZIL: 4 females, OSUC 322670, 348261, 371852, 376063 (OSUC). COLOMBIA: 7 females, OSUC 191148, 193587 (IAVH); OSUC 178096, 191040, 259754, 262600, 276046 (OSUC). ECUADOR: 15 females, OSUC 458510, 458533, 534249–534251, 553237, 553242–553243, 553246, 553406– 553407, 553530, 553651, 553687 (CNCI); OSUC 534659 (OSUC). FRENCH GUIANA: 10 females, OSUC 458384, 458413, 458421, 458459–458460, 458474, 546106–546108, 546135 (CNCI). GRENADA: 6 females, OSUC 534255–534260 (CNCI). PERU: 5 females, OSUC 553952, 553955, 553967, 554016, 554021 (CNCI). TRINIDAD AND TOBAGO: 3 females, OSUC 534601, 534605–534606 (CNCI).

Calliscelio armilaChen & Masner, new species http://zoobank.org/6AB94D9B-DEF6-4FE0-BC1B-5F5BAB7A15F1 http://bioguid.osu.edu/xbiod_concepts/362054 Figures 1.7A–1.7F.

Description. Body length of female: 2.12–2.51 mm (n=11). Body length of male: 2.05–2.27 mm (n=13). Color of head: black throughout; dark brown. Color of antennal clava (A7–A12): black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS slightly less than EH. Sculpture of ventrolateral frons: smooth to rugulose. Sculpture of frons below median ocellus: largely smooth with sparse fine punctures.

37 Sculpture of posterior vertex: coriaceous. Hyperoccipital carina: absent. Occipital carina medially: weakly developed, irregularly sculptured. Length of OOL: greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: coriaceous. Ocular setae: sparse, short. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Form of male antennal flagellomeres: filiform, A11 approximately 3.0x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: variably orange to pale brown. Color of mesosoma in male: orange throughout; variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: largely smooth, granulate ventrally and posteriorly. Sculpture of netrion: rugose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin straight, approximately 3.5x wider than long. Sculpture of metascutellum in female: rugose anteriorly, smooth posteriorly. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugulose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: coxae to femur white, remainder of the legs pale yellow. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: orange to pale brown. Color of metasoma in male: T3–T4 orange, otherwise brown. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: smooth to rugulose medially, with V-shaped keels laterally. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: present on the anterior margin of T2. Sculpture of T3: smooth. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures.

38 Diagnosis. This species is most similar to C. latifrons in the female T1 horn, size and habitus but can be easily distinguished by its smooth upper frons and T3. Etymology. The specific epithet is to be treated as a noun in apposition, derived from the Latin for "bracelet" and refers to the rim on T1 horn. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362054] Material Examined. Holotype, female: HAITI: Sud Dept., Hotte Mts., S slope of Mt. Formon, 31km NW Les Cayes, disturbed forest & fields, 1405m, 18°20'N 74°01'W, Formond, 7.IX–8.IX.1995, R. Davidson, G. Onore & J. Rawlins, OSUC 458342 (deposited in CNCI). Paratypes: HAITI: 10 females, 13 males, OSUC 458334–458341, 458343–458357 (CNCI).

Calliscelio bidensChen & Masner, new species http://zoobank.org/0200D54C-C0FF-41DC-9D85-829CCD8BF6AE http://bioguid.osu.edu/xbiod_concepts/362052 Figures 1.8A–1.8F.

Description. Body length of female: 3.35 mm (n=1). Color of head: brown throughout. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: present. Setation of upper frons: with dense, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth with sparse punctures. Sculpture of frons below median ocellus: granulate to finely punctate. Sculpture of posterior vertex: punctate rugose. Hyperoccipital carina: absent. Occipital carina medially: complete, strongly crenulate throughout. Length of OOL: greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: sparse, short. A4 in female: as long as A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: distinctly longer than wide. Color of mesosoma in female: dark brown. Sculpture of dorsal pronotal area: areolate. Sculpture of lateral pronotal area: smooth dorsally, rugulose ventrally. Sculpture of netrion: rugulose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: densely punctate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: densely punctate. Shape of metascutellum: posterior marging rounded, approximately 4.0x wider than long. Sculpture of metascutellum in female: rugose. Dorsal propodeum in female: not excavate medially, lateral

39 propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose. Median keels on propodeum in female: present. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange yellow. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: dark brown. Horn on T1 in female: absent. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. rugicoxa with which it shares the characters of the hairy compound eyes and the median keels on propodeum in female. It can be distinguished by its smooth hind coxa and the absence of a T1 horn in the female. Etymology. The specific epithet means "with two teeth," refering to the median keels of propodeum of the species and should be treated as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362052] Material Examined. Holotype, female: COLOMBIA: Valle del Cauca Dept., 650m, 03°26'N 76°48'W, Farallones de Cali Natural National Park, 8.V–19.VI.2001, Malaise trap, S. Sarria, OSUC 458326 (deposited in CNCI).

Calliscelio bisulcatus (Kieffer) http://zoobank.org/3C2921D8-CDFA-4CDA-86F2-C61304A4730B http://bioguid.osu.edu/xbiod_concepts/4139 Figures 1.9A–1.9F.

Baryconus bisulcatus Kieffer, 1910a: 320 (original description, keyed). Baryconus (Baryconus) bisulcatus Kieffer: Kieffer, 1910b: 84 (subgeneric assignment). Glyptoteleia bisulcata (Kieffer): Kieffer, 1926: 487 (generic transfer, description). Calliscelio bisulcatus (Kieffer): Masner, 1976: 38 (description, generic transfer, type information).

40

Description. Body length of female: 1.75–2.18 mm (n=20). Body length of male: 1.70–2.12 mm (n=20). Color of head: black throughout; dark brown; orange throughout. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth with sparse punctures. Sculpture of frons below median ocellus: largely smooth with sparse fine punctures. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, strongly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate. Ocular setae: dense, long. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Form of male antennal flagellomeres: filiform, A11 approximately 2.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: orange throughout; variably orange to pale brown. Color of mesosoma in male: orange throughout; variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: rugose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: smooth with sparse punctures. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 2.0x wider than long. Sculpture of metascutellum in female: rugose anteriorly, smooth posteriorly. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: largely smooth with a row of foveae along mesopleural carina. Sculpture of ventral metapleural area: smooth dorsally, densely punctate ventrally. Color of legs: orange throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: approximately as long as r-rs.

41 Color of metasoma in female: orange throughout; variably orange to pale brown. Color of metasoma in male: variably orange to pale brown. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: smooth. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially. Sculpture of T3: smooth. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: smooth. Diagnosis. This species is similar to C. punctatifrons in the shape of metascutellum and sculpture of T1 horn in the female but can be distinguished by its largely smooth upper frons, granulate posterior vertex and smaller body size. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=4139] Material Examined. Holotype, female, B. bisulcatus: BRAZIL: PA, no date, Baker, CAS Type No. 9590 (deposited in CASC). Other material: (216 females, 71 males) ARGENTINA: 8 females, 8 males, OSUC 534117–534124, 534436, 534440–534442 (CNCI); OSUC 63157 (OSUC); OSUC 577254, 577258–577259 (UCRC). BELIZE: 14 females, 1 male, OSUC 534192, 534199–534200, 534202, 534206–534210, 534302–534307 (CNCI). BOLIVIA: 6 females, 2 males, OSUC 534027–534028, 534044, 534049–534053 (CNCI). BRAZIL: 43 females, 23 males, OSUC 534521–534522, 557242, 557297, 557299, 557309–557310 (CNCI); OSUC 110142, 110175, 110182, 111340, 111379, 111382, 111522, 111527, 111531, 111633, 111693–111694, 111697, 111701, 111712–111713, 111920, 111922, 111939, 112040, 112070, 112090, 112186, 112453, 112781, 112806, 113027, 12276, 12356, 130739, 130748, 131475, 132140, 132282, 132534, 132690–132691, 132717, 133048, 133057, 133124, 134453, 134484, 137972, 138060, 232036, 374721–374722, 374726–374727, 48518, 48526, 48548, 48563, 48565, 48568, 48573, 55927, 813 (OSUC). COLOMBIA: 2 females, 3 males, OSUC 557536, 557576, 557624–557625 (CNCI); OSUC 369977 (OSUC). COSTA RICA: 10 females, 1 male, OSUC 532493, 532618–532622, 532646, 532718–532719, 532722, 532740 (CNCI). ECUADOR: 34 females, 10 males, OSUC 458496, 458498, 458500, 458503, 458505–458506, 458508, 458515, 458529, 534229, 534235, 534239–534240, 534242, 553262, 553386–553387, 553389, 553399, 553408–553410, 553497–553499, 553551, 553559, 553594–553595, 553622, 553625, 553634–553636, 553649–553650, 553681, 577329–577330, 577335 (CNCI); OSUC 534658, 534660–534661, 534672 (OSUC). FRENCH GUIANA: 15 females, OSUC 458392, 458398, 546112, 546119, 546122, 546127–546128, 546136–546138, 546140–546141, 546144–

42 546145, 546148 (CNCI). MEXICO: 2 females, OSUC 534459 (CNCI); OSUC 377895 (OSUC). PANAMA: 3 females, OSUC 534083, 534085, 553877 (CNCI). PARAGUAY: 7 females, 18 males, OSUC 185325, 276696, 276729–276731, 276901–276907, 280295, 323025–323026, 323054–323055, 323067, 577193–577194, 577341, 577354, 583310–583311, 583314 (OSUC). PERU: 14 females, 1 male, OSUC 534389 (CNCI); OSUC 323933–323937, 323940–323943, 323945, 323947, 323949–323951 (OSUC). SURINAME: 19 females, OSUC 534562–534567, 534570–534571, 534573–534579, 534582, 553631–553633 (CNCI). TRINIDAD AND TOBAGO: 22 females, OSUC 545999, 546002–546003, 546013, 546027–546028, 546050– 546052, 546057, 546077, 546082, 546086, 546088–546089, 546093–546096, 546099–546100, 553676 (CNCI). URUGUAY: 5 females, OSUC 534610–534614 (CNCI). VENEZUELA: 12 females, 4 males, OSUC 545881, 545884, 545887, 545893, 545895, 545897–545898, 545942– 545943, 557648–557649, 557651–557654 (CNCI); OSUC 55923 (OSUC).

Calliscelio brachysChen & Johnson, new species http://zoobank.org/D4F29FBA-909B-4DE6-969B-BBA983996BD5 http://bioguid.osu.edu/xbiod_concepts/384702 Figures 1.10A–1.10F.

Description. Body length of female: 1.52–1.85 mm (n=20). Body length of male: 1.45–1.88 mm (n=20). Color of head: orange throughout. Color of antennal clava (A7–A12): A7–A9 brown, A10–A12 yellow. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth to coriaceous. Sculpture of frons below median ocellus: coriaceous. Sculpture of posterior vertex: smooth; smooth to transversely striate. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: as long as A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: as long as wide. Form of male antennal flagellomeres: filiform, A11 approximately 3.5x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5.

43 Color of mesosoma in female: orange throughout; orange to pale brown. Color of mesosoma in male: orange throughout; variably orange to pale brown. Sculpture of dorsal pronotal area: smooth. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: abbreviated, at most reaching middle of mesoscutum. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin rounded, approximately 2.5x wider than long. Sculpture of metascutellum in female: rugulose with a longitudinal, median carina. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral to median keel. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth with an oblique carina. Color of legs: orange throughout; pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: greater than 3.0x length of r- rs. Color of metasoma in female: orange throughout; variably orange to pale brown. Color of metasoma in male: variably orange to pale brown. Horn on T1 in female: absent. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: present on the anterior margin of T2. Sculpture of T3: smooth. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: smooth. Diagnosis. This species shares the character of the abbreviated notaulus with C. absum and C. brevinotaulus. It can be separated from C. absum by having A4 as long as A3 in the female and the length of the A5 tyloid in the male is approximately 0.3x length of A5; from C. brevinotaulus it may be distinguished by the absence of the T1 horn in the female. Etymology. The epithet is derived the Greek word for short, in reference to the abbreviated notaulus, and is intended to be treated as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384702]

44 Material Examined. Holotype, female: COLOMBIA: Amazonas Dept., Amacayacu Natural N.P., M.842, 150m, 03°46'S 70°18'W, San Martín, 16.VIII–24.VIII.2000, Malaise trap, B. Amado, OSUC 193349 (deposited in IAVH). Paratypes: (74 females, 26 males) BOLIVIA: 1 female, OSUC 534149 (CNCI). COLOMBIA: 63 females, 24 males, OSUC 557548, 557582 (CNCI); OSUC 152149-152150, 152152, 152154, 182229, 182592, 188678, 188682, 188955, 189186, 189190-189191, 189193, 189195, 189226, 193282, 193284, 193541, 193570, 193584, 193597, 193903, 231811, 249895, 253463, 259760, 259762, 272083, 272087, 275804-275805, 276184, 276236, 276244-276245, 279657, 363591, 364075 (IAVH); OSUC 152162, 162499, 162501, 162509, 162584, 162586-162587, 162594, 162596, 162599, 162606-162607, 162609, 170495, 182596, 182722-182723, 182740, 188551, 188941, 188944, 188963, 189183, 189267- 189268, 189271, 189273, 189279, 191094, 191150, 192353, 192358, 193320-193321, 193329, 193338, 193340, 193538, 193598-193599, 193814, 231829, 267807, 267962, 280198, 372644- 372645 (OSUC). ECUADOR: 5 females, 1 male, OSUC 458488, 458497, 458536, 534237, 553561, 577338 (CNCI). PERU: 5 females, 1 male, OSUC 553970, 553972, 554010, 554023– 554024, 554032 (CNCI).

Calliscelio brevinotaulusChen & Johnson, new species http://zoobank.org/9D986838-195F-44DD-BF84-3D933502679D http://bioguid.osu.edu/xbiod_concepts/363559 Figures 1.11A–1.11F.

Description. Body length of female: 1.78–2.98 mm (n=20). Body length of male: 1.78–2.41 mm (n=20). Color of head: black throughout; variably orange to dark brown. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: transversely rugulose to granulate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: largely smooth with small granulate area. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape

45 of female A6: subquadrate. Form of male antennal flagellomeres: filiform, approximately 2.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: orange throughout; variably orange to pale brown. Color of mesosoma in male: orange throughout; variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout; smooth anteriorly, granulate posteriorly. Sculpture of netrion: rugose. Notaulus: abbreviated, at most reaching middle of mesoscutum. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin rounded, 2.5x wider than long. Sculpture of metascutellum in female: rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral to median keel. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange throughout; mid and hind coxae dark brown to black, otherwise yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: greater than 3.0x length of r- rs. Color of metasoma in female: orange throughout; variably orange to pale brown. Color of metasoma in male: orange throughout; variably orange to pale brown. Horn on T1 in female: weakly indicated. Sculpture of T1 horn dorsally: with V-shaped striae. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth; largely smooth with submedian longitudinal striae. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. This species is most similar to C. brachys in color, size and the abbreviated notaulus but can be distinguished by the granulate upper frons and that A4 is shorter than A3.

46 Etymology. The epithet refers to the short notauli in this species and is intended to be used as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=363559] Material Examined. Holotype, female: COLOMBIA: Magdalena Dept., Zaino, M.564, 50m, 11°20'N 74°02'W, Tayrona Natural National Park, 14.VIII–30.VIII.2000, Malaise trap, R. Henriquez, OSUC 190678 (deposited in IAVH). Paratypes: (264 females, 99 males) BELIZE: 7 females, 5 males, OSUC 534277, 534279, 534283–534284, 534290–534292, 534301, 534308, 534310–534311, 534313 (CNCI). BRAZIL: 29 females, 1 male, OSUC 348007, 348118, 353716, 363846, 366704-366705, 371850, 374564, 374566, 374593, 376520-376521, 380186, 577137 (MZSP); OSUC 326244, 347202, 347274, 347666, 347738, 347969, 348098, 349378, 349773, 351396, 351679, 351831-351832, 351839-351840, 351843 (OSUC). COLOMBIA: 70 females, 61 males, OSUC 557546, 557554, 557559, 557561, 557569-557573, 557584, 557621, 557628, 557634-557639 (CNCI); OSUC 170449-170452, 170497, 170503-170504, 170506, 188889, 189906, 189910, 190095, 190097-190098, 190100, 190124, 190706-190707, 190709, 190986, 190988-190989, 190992, 190994, 191057, 191061, 191143, 191739, 191742-191743, 191745, 192204, 192211, 192214, 193379, 193701, 193706-193709, 193772, 259573, 259583, 269666, 273785, 364080-364086, 370058, 370060, 370063-370064 (IAVH); OSUC 188887, 188890, 189390-189391, 189396, 189398, 190666, 190672-190673, 190677, 190680-190681, 190683, 191064, 191241-191242, 191245, 191247, 191249-191250, 191253-191258, 191268, 191272-191276, 191278, 192202, 192215-192216, 192385-192390, 192394, 193704, 193778, 194185, 194189, 194194, 194200, 259572, 259574, 259577, 259579, 259582, 259588-259589, 279454 (OSUC). COSTA RICA: 52 females, 22 males, OSUC 532464, 532479–532486, 532506, 532516, 532518–532519, 532523–532526, 532532, 532565, 532572–532574, 532577– 532578, 532583, 532585–532588, 532590–532591, 532602–532608, 532611, 532632–532634, 532643, 532665–532666, 532675–532676, 532678–532681, 532685, 532687, 532698–532700, 532717, 532720–532721, 532732, 532734–532735, 532737–532739, 532751–532752, 532756– 532757, 532788, 532790, 532834, 532919 (CNCI); OSUC 237329 (OSUC). CURAÇAO: 2 males, OSUC 532749–532750 (CNCI). ECUADOR: 30 females, OSUC 458501–458502, 458532, 458543–458544, 534228, 553383–553384, 553396, 553398, 553404, 553417, 553483, 553500, 553537, 553548, 553558, 553591, 553593, 553626, 553629, 553637–553639, 553679, 553683, 553710, 553712, 553737, 553746 (CNCI). HONDURAS: 4 females, OSUC 534142–

47 534144, 534146 (CNCI). MEXICO: 19 females, 4 males, OSUC 533982–533997, 534001– 534003, 534025, 534460, 534488–534489 (CNCI). PANAMA: 26 females, 2 males, OSUC 534066–534068, 534082, 534084, 534101–534102, 553773–553775, 553808–553809, 553814, 553816, 553820, 553872, 553904, 553919, 553926–553927, 553938–553940, 553944–553947 (CNCI); OSUC 55955 (OSUC). PERU: 1 female, OSUC 554025 (CNCI). SURINAME: 1 female, OSUC 534585 (CNCI). VENEZUELA: 25 females, 2 males, OSUC 532724, 545833– 545834, 545847–545848, 545851, 545864–545868, 545879–545880, 545899, 545958–545959, 545968–545973, 557656, 557658, 557673 (CNCI); OSUC 334558, 360696 (OSUC).

Calliscelio brevitasChen & Johnson, new species http://zoobank.org/10B4E8E2-0970-4E2A-85EC-7B373111AFDF http://bioguid.osu.edu/xbiod_concepts/367290 Figures 1.12A–1.12F.

Description. Body length of female: 1.83–2.88 mm (n=20). Color of head: black throughout; orange throughout; variably orange to dark brown. Color of antennal clava (A7–A12): A7 dark orange, remainder dark brown to black; dark brown to black. Shape of head: subglobose. Central keel of frons: present. Setation of upper frons: with sparse, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate; smooth with sparse punctures. Sculpture of frons below median ocellus: largely smooth with sparse fine punctures. Sculpture of posterior vertex: densely punctate; granulate to rugulose. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: as long as wide. Color of mesosoma in female: orange throughout; black throughout; variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout; largely smooth, granulate ventrally and posteriorly. Sculpture of netrion: smooth; rugose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: largely coriaceous with dense and fine punctures at poster end. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of

48 mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth; largely smooth, rugose ventrally. Color of legs: pale yellow throughout; coxae pale yellow, otherwise orange to pale brown. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: variably yellow to pale brown. Color of metasoma in male: brown throughout; reddish brown. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: transversely striate. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2; present on anterior margin of T2 medially, reaching posterior margin of T2 laterally. Sculpture of T3: smooth with longitudinal submedian striae. Shape of T6 in female: distinctly elongate, at least 2.0x longer than wide. Sculpture of S3: densely punctate. Diagnosis. This species is most similar to C. laticinctus and C. longius in color, size and habitus. It may be distinguished from them by having A6 quadrate in female, whereas A6 is distinctly longer than wide in C. laticinctus and C. longius. Etymology. The epithet is used as a noun in apposition derived from the Latin word for shortness, in reference to the shortened A5 and A6. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=367290] Material Examined. Holotype, female: COLOMBIA: Magdalena Dept., Zaino, M.567, 50m, 11°20'N 74°02'W, Tayrona Natural National Park, 28.VII–14.VIII.2000, Malaise trap, R. Henriquez, OSUC 190705 (deposited in IAVH). Paratypes: (355 females) BRAZIL: 11 females, OSUC 534537, 534539 (CNCI); OSUC 121827, 252086, 323928, 326208, 348271, 349206, 355169, 376519, 376528 (OSUC). COLOMBIA: 126 females, OSUC 557562-557564, 557568,

49 557581, 557583, 557590, 557592-557597, 557603, 557626-557627 (CNCI); OSUC 144164, 144230, 162601, 166586, 170369-170371, 178098, 178162, 178174, 178192-178193, 178195, 190125, 191183, 191206, 191212, 191243, 191248, 191381, 191383, 193128, 193130, 193179, 193193, 193294, 193322, 193326, 193328, 193863-193864, 193866-193868, 193879, 193906, 193910, 193913, 262616, 267805, 363594, 76997 (IAVH); OSUC 178021, 178191, 182228, 182482, 182754-182756, 182762, 188688, 188726, 188730, 188951, 188954, 189176, 189199, 189277, 189282, 189285, 189287, 189291-189292, 189297, 190310, 191096, 191098, 191827, 192354-192355, 192359, 193166, 193293, 193560, 193563, 193565-193566, 193572, 193576, 193689-193691, 194196, 202078, 232297-232298, 253454-253455, 268911, 269217, 269350, 269356, 275800, 275803, 275808, 275811, 279350, 279899, 279903, 279908-279909, 279911, 280179, 280181, 280195, 280204-280205, 363600, 374718, 377424 (OSUC). COSTA RICA: 133 females, OSUC 532458, 532503, 532535, 532570, 532616–532617, 532627–532628, 532630–532631, 532635, 532637–532639, 532660, 532771, 532774, 532784–532785, 532825, 532841–532845, 532847–532848, 532850, 532852–532868, 532870–532872, 532875–532885, 532887–532888, 532891–532892, 532899–532901, 532903–532916, 532928, 532935, 532938– 532940, 532942–532949, 534138, 557113–557132, 557134–557139, 557142–557154 (CNCI). ECUADOR: 29 females, OSUC 458494, 553255, 553376, 553440, 553443, 553461, 553469– 553471, 553473, 553478, 553510–553511, 553513–553514, 553518, 553524–553526, 553540, 553542, 553565, 553568–553570, 553590 (CNCI); OSUC 534663, 534668, 534670 (OSUC). FRENCH GUIANA: 25 females, OSUC 458393–458397, 458404–458405, 458410–458411, 458423, 458427, 458444–458446, 458462, 458464, 458469, 458472–458473, 458478, 458480– 458481, 546104, 546132, 546149 (CNCI). PANAMA: 17 females, OSUC 534100, 553747, 553756, 553767–553770, 553782, 553784, 553810, 553905, 553915, 553924–553925, 553930– 553932 (CNCI). PERU: 12 females, OSUC 534416, 553991, 553993, 554035, 554043, 554047, 554049 (CNCI); OSUC 570539–570540, 570546 (OSUC); OSUC 228135, 228191 (USNM). SURINAME: 1 female, OSUC 534583 (CNCI). VENEZUELA: 1 female, OSUC 557696 (CNCI).

Calliscelio carinigenaChen & Johnson, new species http://zoobank.org/ADB95787-EBBB-4A36-B75D-5CE10D93826F http://bioguid.osu.edu/xbiod_concepts/363281

50 Figures 1.13A–1.13F.

Description. Body length of female: 1.70–2.46 mm (n=20). Body length of male: 2.00–2.29 mm (n=20). Color of head: black throughout; orange throughout; variably brown to black. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth to granulate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate to rugulose above hyperoccipital carina, smooth below. Hyperoccipital carina: present. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: with a carina along outer orbit. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Form of male antennal flagellomeres: filiform, A11 approximately 3.0x longer than wide. Length of A5 tyloid in male: longer than 0.5x length of A5. Color of mesosoma in female: orange throughout. Color of mesosoma in male: orange throughout; variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin rounded, 2.5x wider than long. Sculpture of metascutellum in female: rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange throughout; pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: greater than 3.0x length of r- rs.

51 Color of metasoma in female: orange throughout; yellow throughout; T1 horn and T6 pale brown, otherwise orange throughout. Color of metasoma in male: orange throughout; variably orange to pale brown. Horn on T1 in female: weakly developed. Sculpture of T1 horn dorsally: smooth to somewhat transversely striate. Sculpture of posterior margin of T1 in female: smooth medially, longitudinally striate laterally. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially. Sculpture of T3: smooth. Shape of T6 in female: short, wider than long. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. crater and C. sfina. It can be separated from C. crater by the presence of a carina along the outer orbit on the postgena rather than large foveae, and from C. sfina by the presence of the hyperoccipital carina. Etymology. The epithet is a compound noun in reference to the genal carina along the outer orbit. It is intended to be used as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=363281] Material Examined. Holotype, female: BRAZIL: SP, Pilões Trail, Boracéia Biological Station, pt.4, 23°39'05.0"S 45°53'40.1"W, Salesópolis, 19.X–22.X.2002, yellow pan trap, A. P. Aguiar & J. S. Freitas, OSUC 343907 (deposited in MZSP). Paratypes: (941 females, 591 males) ARGENTINA: 1 female, OSUC 534443 (CNCI). BOLIVIA: 4 females, 1 male, OSUC 534041–534043, 534056, 534064 (CNCI). BRAZIL: 929 females, 582 males, OSUC 557167- 557179, 557182-557187, 557189-557196, 557210-557219, 557221-557241, 557244-557245, 557247, 557249-557253, 557256-557281, 557284-557295, 557300-557308, 557311-557317, 557322-557324, 557326-557336, 557339, 557341, 557343-557348, 557350-557359, 557363- 557374, 557378-557394, 557396-557408 (CNCI); OSUC 577148, 577309-577311, 577313, 577379-577381, 577384-577387, 577389 (MNHN); OSUC 135930, 135955, 136029, 136037, 136420, 136453, 136953, 136964, 137063, 137244, 137263, 137306, 137403, 137408, 137429, 137694-137695, 137702, 137711-137712, 137740, 137743, 137753, 137762, 137766, 137768, 137771, 137775, 137777, 137806, 137890, 137894, 137912, 137916, 137927, 138530, 138536, 138542-138543, 138546, 138693, 138699-138701, 138721-138722, 138724, 138894, 138987, 139006, 139016, 139029, 150129-150152, 150154-150155, 150157, 150208-150250, 150265- 150279, 150281-150285, 150357, 150371-150372, 150393-150394, 150419, 237725-237726, 318263-318266, 318268-318276, 318278-318280, 318961-318963, 319014-319021, 319023- 319027, 319080-319139, 319340-319348, 319390-319400, 320901, 322525-322527, 322529-

52 322531, 322541-322543, 322550-322551, 323094-323098, 323286, 323308, 323324, 323328, 323336, 343702, 343899-343906, 343908-343913, 343915-343931, 345915-345917, 345919, 345922, 345925, 345938, 345945, 345955, 345958-345959, 345962-345965, 345967-345968, 345970-345978, 345980-345983, 345985-345988, 345991, 345997, 346002, 346004-346006, 346011-346012, 346021, 346056-346057, 346059, 346062, 346068, 346072-346075, 346095, 346097-346099, 346112-346113, 349591, 349593, 352850-352856, 352859-352861, 353270- 353272, 353870-353873, 356331, 356946-356947, 356949-356950, 356967-356969, 356973, 356975, 356982, 357249, 357251, 357253-357254, 357259-357261, 357264, 357319, 363768- 363773, 363880, 366659, 371817-371823, 371825-371827, 372536, 372539, 373870-373872, 374591, 374599-374621, 375255, 375261, 375288, 375290, 375293, 375296, 376556, 376921, 376959, 377461-377465, 377467-377480, 377933-377959, 377961-377968, 377970-377977, 378977, 378982-378984, 381077-381078, 381080-381085, 433842, 433844, 433852, 576982, 577005-577013, 577034-577035, 577048, 577058-577059, 577064-577066, 577071, 577097, 577100, 577113-577115, 577122-577124, 577217, 577221-577223, 577228-577229, 577237- 577245, 577262-577267, 577269-577271, 577277-577278, 577281-577282, 577284-577286, 577289, 577294, 583206-583208, 583213-583225, 583233, 583235, 583240, 583242, 583259- 583265, 583269, 583281-583288, 583293-583295, 583297 (MZSP); OSUC 135931, 135938, 135952-135953, 135966, 135979, 135989-135990, 136007, 136009, 136125, 136128, 136135, 136190, 136192, 136195, 136217, 136225, 136242, 136364-136365, 136368, 136370, 136373, 136375, 136379-136380, 136392, 136423, 136450, 136511-136512, 136519, 136526, 136529, 136535, 136547-136548, 136550, 136560, 136567, 136571, 136618, 136628, 136638, 136646, 136648, 136659, 136669, 136696, 136705, 136709, 136735-136736, 136760-136761, 136785, 136834, 136839-136840, 136843, 136849, 136852, 136858, 136864-136865, 136874, 136881, 136883, 136892, 136906, 136910, 136913, 136919, 136928, 136933, 136950-136951, 136959, 136962, 136967, 136969-136970, 136972, 136983, 136986, 136998, 137005, 137008, 137010, 137013, 137017-137018, 137027, 137054-137055, 137059, 137068-137069, 137075, 137079, 137084, 137119, 137134, 137173-137174, 137178, 137180, 137185, 137188-137189, 137200, 137206, 137212-137213, 137223-137224, 137227, 137230, 137234-137235, 137238, 137241, 137246, 137248, 137250, 137256, 137311, 137314, 137336, 137342, 137344-137345, 137349, 137354, 137357, 137361-137362, 137368-137370, 137372, 137379, 137384, 137393, 137398- 137401, 137409, 137413, 137419-137423, 137428, 137434-137435, 137438, 137441-137442,

53 137448, 137455, 137507, 137509, 137523, 137525, 137531, 137533-137535, 137543, 137547- 137549, 137551, 137555, 137564-137565, 137573, 137576, 137581, 137587, 137591, 137606- 137609, 137646, 137649-137650, 137653-137654, 137656, 137660-137661, 137663-137664, 137668, 137685, 137718, 137729-137730, 137738, 137745, 137788, 137796, 137809, 137822, 137834, 137840, 137849, 137852-137853, 137861, 137863, 137865, 137873, 137880, 137886- 137887, 137889, 137899, 138421, 138426, 138431-138432, 138434-138435, 138438, 138441- 138445, 138448-138449, 138452, 138455, 138460-138462, 138465, 138468-138469, 138471, 138473, 138478, 138484, 138489, 138508-138509, 138515, 138520-138521, 138525, 138553- 138554, 138560, 138562, 138566-138568, 138573, 138575, 138583, 138585-138586, 138593, 138597, 138613-138615, 138622, 138627, 138631-138632, 138636-138637, 138639, 138641, 138643, 138647, 138651, 138669-138670, 138677-138678, 138680, 138682, 138685, 138688, 138691-138692, 138705, 138710, 138713, 138715, 138725, 138737, 138739, 138743, 138746, 138749-138750, 138753, 138755-138757, 138760-138761, 138770-138771, 138774, 138776, 138783, 138786, 138788, 138791, 138805, 138808, 138813, 138815, 138819, 138821-138822, 138824, 138828, 138833, 138841, 138846, 138857, 138873, 138876, 138878, 138880, 138889- 138890, 138892, 138895-138896, 138898, 138900, 138906, 138922, 138925, 138942, 138955, 138958, 138982, 139013, 139015, 139017, 139021, 139065, 139067, 139079-139080, 139089, 139091, 139093-139094, 139096, 139102, 139106, 139109, 139113, 139117-139119, 139121, 139130-139131, 139133, 139136-139137, 139143, 139146-139147, 139152, 139157-139159, 139164-139166, 139168, 139172, 139175, 139179-139180, 150158, 150160-150163, 150696- 150700, 150736-150737, 150743, 150964, 150985, 151153-151154, 151163, 151189-151193, 151216-151223, 151225-151226, 151247-151252, 151254, 151267, 151270-151271, 151290- 151293, 151309, 151344-151345, 151368-151373, 151384, 151386, 151398-151399, 151410, 151452-151480, 151488, 151492-151494, 151498, 151506, 151517-151519, 151526, 151593, 151629-151632, 319022, 322988, 323099, 323319, 323526, 334296-334298, 343914, 346078, 352843-352845, 352847-352849, 359023-359025, 365999-366000, 366656, 367439, 377481, 377960, 378585, 378591-378592, 40024, 40036, 40046, 40061, 40071, 40099, 40164, 40175, 40303, 42295, 42306-42309, 42312, 42315, 42320, 42328-42330, 42334, 42338, 42342-42343, 42354, 42361, 42363-42364, 42367-42368, 42375, 42378-42379, 427454, 433823, 433832, 433837, 463268, 463307-463308, 510886, 510888, 510893, 55909-55921, 55957-55997, 577015-577016, 577018, 577020, 583270-583280, 583298-583304, 583306-583307, 58793,

54 61197, 61236, 62250, 62261, 62268, 62284, 62287, 62301, 62312, 62333, 62348 (OSUC). COLOMBIA: 3 males, OSUC 162611, 191322, 191324 (OSUC). ECUADOR: 4 males, OSUC 534673–534675, 534679 (OSUC). PARAGUAY: 1 female, 1 male, OSUC 278829, 577183 (OSUC). TRINIDAD AND TOBAGO: 2 females, OSUC 546058, 546060 (CNCI). VENEZUELA: 4 females, OSUC 545888, 545891–545892, 557655 (CNCI).

Calliscelio craterChen & Johnson, new species http://zoobank.org/3643BD0B-9BF3-4C41-A861-2AB72C701CA3 http://bioguid.osu.edu/xbiod_concepts/363278 Figures 1.14A–1.14F.

Description. Body length of female: 1.74–2.35 mm (n=20). Body length of male: 1.86–2.36 mm (n=20). Color of head: black throughout; variably orange to dark brown. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth to granulate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate above hyperoccipital carina, smooth to rugulose below. Hyperoccipital carina: present. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: with large foveae. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: subquadrate. Form of male antennal flagellomeres: filiform, A11 approximately 2.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: orange throughout. Color of mesosoma in male: orange throughout; variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth anteriorly, granulate posteriorly. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin straight, approximately 2.5x wider than long. Sculpture of metascutellum in female: rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: shallowly

55 excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: orange throughout; T1 horn and T6 pale brown, otherwise orange throughout. Color of metasoma in male: variably orange to pale brown. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: rugose medially, with V-shaped keels laterally. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially. Sculpture of T3: coriaceous; granulate; smooth medially, coriaceous laterally; longitudinally striate throughout. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: smooth to coriaceous. Diagnosis. This species is similar to C. carinigena and C. sfina in color, size and habitus, but it can be easily recognized by the large foveae on the postgena (postgena of C. sfina granulate, whereas C. carinigena with a carina along outer orbit). Etymology. The specific epithet refers to the foveae on the postgena and should be treated as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=363278] Material Examined. Holotype, female: PARAGUAY: Guairá Dept., Amambay Stream, Pa'i Thermal Waterfall, second growth forest, 221m, 25°44'56"S 56°15'12"W, Melgarejo, 24.III– 25.III.2005, yellow pan trap, Garcete, OSUC 276803 (deposited in MNHNPY). Paratypes: (228 females, 322 males) BOLIVIA: 8 females, OSUC 534035–534037, 534039, 534057, 534059, 534161, 534165 (CNCI). BRAZIL: 202 females, 276 males, OSUC 534519, 534524, 534546- 534547, 534621 (CNCI); OSUC 577298, 577305 (MNHN); OSUC 10431, 10463, 10584, 10625, 10833, 10857, 110188, 110198, 131683, 134141, 134350, 134445, 134795, 134802, 134829, 134845, 135140, 135614, 135652-135653, 137956, 137964, 137982, 138015, 138041, 138116, 138181, 147790, 151160, 151162, 232033, 232035, 323264-323266, 323269-323271, 323273-

56 323274, 323276-323279, 323281, 323283-323285, 323288-323291, 323296-323297, 323299- 323301, 323303-323304, 323311-323316, 323318, 323320, 323322, 323326, 323332-323335, 323525, 323529-323530, 323533-323537, 323539-323541, 323543, 323545, 323548, 323550, 323553-323557, 323561-323562, 323564-323565, 323914, 323918, 356945, 357216-357223, 357226-357228, 357257-357258, 357262-357263, 357273, 357278, 358790, 358944, 358984, 375277, 375285-375287, 375295, 375300, 376550-376551, 376553, 376555, 376947-376954, 376956-376958, 376960-376961, 376964, 433819, 510906, 534641-534642, 534647, 577019, 577022-577031, 577036, 577039-577042, 577044-577045, 577047, 577049-577057, 577062, 577091-577093, 577095, 577098, 577102, 577109-577112, 577198, 577205-577210, 577212- 577214, 577218-577219, 577230-577233, 577235-577236, 577249-577252, 577268, 577276, 577279-577280, 577283, 577288, 577290-577292, 577295, 583228-583229, 583234, 583237- 583239, 583241, 583243-583245, 583267-583268 (MZSP); OSUC 10135, 10363, 10467, 10473, 10523-10524, 10527, 10572-10573, 10581, 10702, 10712, 12201, 12262, 12266, 12325, 131326, 131681, 131720, 131771, 131825, 131923, 131942, 131997, 134140, 134185, 134523, 134565, 134682, 13550, 137264, 137987, 138000, 138479, 13902, 150741-150742, 150744, 150746- 150747, 150749-150750, 150956, 150959, 150974, 150984, 151097, 151152, 151155-151158, 151161, 151266, 151268-151269, 151346, 151383, 151385, 151387, 151401-151404, 151416- 151423, 151485-151487, 151549, 159, 187, 206, 216, 323511-323512, 323514, 323516, 323523- 323524, 323912, 323919, 347618-347619, 349106, 357241-357242, 357247-357248, 357250, 357252, 357312-357313, 357315-357318, 357320, 357323, 357325, 358786-358788, 358942, 358974, 358981-358983, 358992, 363867, 363869-363875, 363877-363879, 363883-363887, 363889-363891, 366657, 366665, 372528, 372530-372534, 372538, 372540-372544, 375246, 375248-375250, 375252, 375254, 375256-375260, 375262, 375264-375268, 375270-375276, 375278-375284, 375297, 376552, 376557-376564, 376922, 376924-376929, 376931-376932, 376934, 376936-376940, 376944-376946, 376965, 477163-477164, 48517, 48537, 48543, 48552, 510889-510892, 510894-510898, 510900, 510905, 534643-534645, 534649, 534651- 534656, 539, 577021, 577046, 712 (OSUC). PARAGUAY: 17 females, 46 males, OSUC 348870, 404893-404894, 534698, 534701, 534703, 534705, 534708-534712, 534715-534719, 577175, 577184-577187, 577339, 577348, 577351-577353 (MNHNPY); OSUC 577177, 577181-577182, 577190, 577342 (MZSP); OSUC 150604, 150608-150609, 150612, 185381, 266183, 276794-276795, 276797, 276799-276802, 277313-277314, 278659-278660, 278671-

57 278672, 278680, 278827, 278831, 280439, 323004, 323030-323031, 323033, 323913, 353273, 534729-534730 (OSUC). VENEZUELA: 1 female, OSUC 545855 (CNCI).

Calliscelio crenaChen & Johnson, new species http://zoobank.org/C53F1631-C068-4EF9-AB3D-39BA37F7A09F http://bioguid.osu.edu/xbiod_concepts/384634 Figures 1.15A–1.15F.

Description. Body length of female: 2.95–3.51 mm (n=20). Body length of male: 2.24–2.37 mm (n=6). Color of head: black throughout; dark brown. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth with sparse punctures. Sculpture of frons below median ocellus: largely smooth with sparse fine punctures. Sculpture of posterior vertex: largely smooth with sparse fine punctures. Hyperoccipital carina: absent. Occipital carina medially: complete, strongly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: as long as wide. Form of male antennal flagellomeres: filiform, A11 approximately 2.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: brown. Color of mesosoma in male: brown throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth anteriorly, granulate posteriorly. Sculpture of netrion: rugulose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: anterior margin rugulose, remainder smooth. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral to

58 median keel. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange throughout; coxae to femurs pale yellow, otherwise orange. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: dark brown. Color of metasoma in male: brown throughout. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: transversely striate. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth with longitudinal submedian striae. Shape of T6 in female: distinctly elongate, approximately 3.0x longer than wide. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. brevitas in color, size and habitus but can be distinguished by its strongly crenulate occipital carina and relatively smooth posterior vertex (C. brevitas with occipital carina weakly crenulate, posterior vertex densely punctate). Etymology. The epithet is used as a noun in apposition derived from the Latin word for notch, in reference to the strongly crenulate occipital carina medially. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384634] Material Examined. Holotype, female: ECUADOR: Santo Domingo de los Tsáchilas Prov., 16km SE Santo Domingo de los Colorados, Tinalandia, 680m, 4.V–25.VII.1985, malaise trap/flight intercept trap, S. Peck & J. Peck, OSUC 553462 (deposited in CNCI). Paratypes: (28 females, 6 males) COLOMBIA: 3 females, OSUC 534555, 557567 (CNCI); OSUC 279904 (OSUC). ECUADOR: 25 females, 6 males, OSUC 534227, 553367, 553373, 553382, 553437– 553439, 553450–553451, 553456, 553464–553466, 553468, 553547, 553600, 553653, 553669, 553675, 553680, 553692–553699, 553705, 553707, 553717 (CNCI).

Calliscelio eborisChen & Johnson, new species http://zoobank.org/71300D0B-D933-4E9B-A689-0C7EA098F418 http://bioguid.osu.edu/xbiod_concepts/384800 Figures 1.16A–1.16F.

59

Description. Body length of female: 1.83–2.27 mm (n=20). Body length of male: 1.94–2.19 mm (n=20). Color of head: variably pale yellow to brown; reddish orange throughout. Color of antennal clava (A7–A12): A7–A9 brown, A10–A12 white; A7 brown, remainder orange. Shape of head: subglobose. Central keel of frons: present. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth to coriaceous. Sculpture of frons below median ocellus: coriaceous. Sculpture of posterior vertex: transversely striate. Hyperoccipital carina: present. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: as long as A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: as long as wide. Form of male antennal flagellomeres: filiform, A11 approximately 3.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: variably yellow to pale brown; reddish orange throughout. Color of mesosoma in male: orange throughout; variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: coriaceous. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin rounded, approximately 3.0x wider than long. Sculpture of metascutellum in female: smooth with a longitudinal, median carina. Sculpture of metascutellum in male: smooth with longitudinal, median carina. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose with one or two longitudinal keels lateral median keel. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral median keel. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange throughout; pale yellow throughout. Sculpture of hind coxa: smooth.

60 Color of fore wing: hyaline. Rs+M: spectral. Setae on R: short, decumbent, hardly exceeding the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: orange throughout; variably orange to pale brown. Color of metasoma in male: variably orange to pale brown. Horn on T1 in female: absent. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially. Sculpture of T3: smooth. Shape of T6 in female: short, wider than long. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. brachys and C. suni. It can be separated from C. brachys by the percurrent notaulus, and from C. suni by the spectral Rs+M and the presence of a central keel on the frons. Etymology. The epithet is used as a noun in apposition derived from the Latin word for ivory, in reference to the white A11–A12 of the female antennae. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384800] Material Examined. Holotype, female: COLOMBIA: Amazonas Dept., Amacayacu Natural N.P., M.84, 150m, 03°41'S 70°15'W, Matamata, 6.IV–15.IV.2000, Malaise trap, A. Parente, OSUC 193404 (deposited in IAVH). Paratypes: (58 females, 26 males) BOLIVIA: 12 females, 5 males, OSUC 534030, 534150–534151, 534156–534157, 534166–534171, 534173– 534176, 534178–534179 (CNCI). COLOMBIA: 30 females, 19 males, OSUC 557578-557580 (CNCI); OSUC 143977, 152163, 152166, 162512, 162608, 176895, 189272, 193278, 193324- 193325, 193327, 193424, 193461, 193932, 269448, 274572, 276241 (IAVH); OSUC 152164, 182227, 182485, 182492-182493, 182496, 182591, 182718-182720, 182733, 188948, 188959, 188962, 189175, 189205, 189216, 189275, 189283, 189290, 190305-190306, 192356, 193583, 193600, 231804, 231821, 272090, 279658 (OSUC). PERU: 11 females, 2 males, OSUC 553953, 553956, 553965, 554055–554064 (CNCI). VENEZUELA: 5 females, OSUC 223877–223881 (USNM).

Calliscelio elegans (Perkins) http://zoobank.org/6B386A5E-E5DF-48CB-A4C9-C82160E262C5 http://bioguid.osu.edu/xbiod_concepts/245756

61

Caloteleia elegans Perkins, 1910: 624 (original description). Caenoteleia elegans (Perkins): Kieffer, 1926: 550 (generic transfer, description). Calotelea tanugatra Narendran, 1998: 71 (original description, keyed); Rajmohana K., 2006: 122, 123 (description, keyed); Rajmohana, Peter & Narendran, 2013: 8 (junior synonym of Calliscelio elegans (Perkins), type information). Calliscelio elegans (Perkins): Masner, Johnson & Musetti, 2009: 61 (description, diagnosis, generic transfer); Rajmohana, Peter & Narendran, 2013: 8 (description of male, synonymy).

Description. See Masner et al. (2009) and Rajmohana et al. (2013). Diagnosis. Callliscelio elegans is easily distinguished within Calliscelio based on the combination of the following characters: fore wing with three darkened and two white bands; head and mesosonotum granulose; metascutellum extremely narrow and weakly concave medially to accommodate T1 horn (Masner et al. 2009). Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=245756] Material Examined. Non-type material: (61 females) AUSTRALIA: 1 female, OSUC 256856 (CNCI). BELIZE: 4 females, OSUC 256871–256874 (CNCI). BENIN: 3 females, OSUC 256882–256884 (CNCI). CHRISTMAS ISLAND: 4 females, OSUC 256857–256860 (ANIC). FIJI: 2 females, FBA015304, OSUC 256864 (BPBM). FRENCH POLYNESIA: 8 females, OSUC 256861–256863 (CNCI); OSUC 256866–256867 (UCDC); UCRC ENT 111562, 135651–135652 (UCRC). GUAM: 1 female, OSUC 256854 (CNCI). INDIA: 3 females, OSUC 256899–256901 (CNCI). INDONESIA: 4 females, OSUC 256893–256896 (CNCI). IVORY COAST: 2 females, OSUC 256885–256886 (CNCI). MADAGASCAR: 2 females, CASENT 2029779, OSUC 215759 (CAS). MAURITIUS: 1 female, MHNG 0002 (MHNG). MEXICO: 5 females, OSUC 256868–256870 (CNCI); OSUC 256875 (NMNH); OSUC 583205 (OSUC). NEPAL: 4 females, OSUC 256890–256892, 256898 (CNCI). NIGERIA: 1 female, OSUC 256881 (CNCI). PUERTO RICO: 2 females, OSUC 256876–256877 (CNCI). SAMOA: 1 female, OSUC 256865 (BPBM). SRI LANKA: 2 females, OSUC 256902–256903 (CNCI). THAILAND: 3 females, OSUC 256897, 256907, 321997 (OSUC). UNITED STATES: 3 females, OSUC 256855 (CNCI); OSUC 256878 (NMNH); UCFC 0 079 680 (UCFC).

62 VENEZUELA: 2 females, OSUC 256879–256880 (CNCI). YEMEN: 3 females, OSUC 256887–256889 (CNCI).

Calliscelio extenuatusChen & Johnson, new species http://zoobank.org/CE5AD0AA-DCC9-4645-AE36-45C93748A3EF http://bioguid.osu.edu/xbiod_concepts/384360 Figures 1.17A–1.17F.

Description. Body length of female: 1.34–1.74 mm (n=20). Body length of male: 1.81 mm (n=1). Color of head: brown throughout; orange throughout; orange to pale brown. Color of antennal clava (A7–A12): dark brown to black; A7, A8 brown, A9–A12 white to pale yellow. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: glabrous. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth to coriaceous; smooth to granulate. Sculpture of frons below median ocellus: smooth; smooth to granulate. Sculpture of posterior vertex: rugose; granulate to rugulose. Hyperoccipital carina: absent. Occipital carina medially: weakly indicated, irregularly sculptured. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: largely smooth with small granulate area. Ocular setae: sparse, short. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Color of mesosoma in female: orange throughout; yellow throughout; orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin rounded, approximately 3.0x wider than long. Sculpture of metascutellum in female: rugose. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange throughout; pale yellow throughout. Sculpture of hind coxa: smooth.

63 Color of fore wing: hyaline. Rs+M: nebulose, weakly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: orange throughout; variably orange to pale brown; yellow throughout. Horn on T1 in female: absent. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially; reaching posterior margin of T2. Sculpture of T3: smooth. Shape of T6 in female: short, wider than long. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. This species is most similar to C. ruga and C. suni in size and habitus. It can be separated from C. ruga by the complete occipital carina and the shortened A6, from C. suni by the hairy compound eyes. Etymology. The epithet is an adjective, the Latin word for faint, weak and reduced, in reference to the weakly indicated occipital carina. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384360] Material Examined. Holotype, female: VENEZUELA: Aragua St., Rancho Grande Biological Station, 1100m, 10°22'N 67°41'W, Henri Pittier National Park, 10.VIII–13.VIII.1994, flight intercept trap, M. Archangelsky, OSUC 48454 (deposited in OSUC). Paratypes: (170 females, 1 male) COLOMBIA: 1 female, OSUC 279960 (OSUC). COSTA RICA: 1 female, OSUC 237330 (OSUC). TRINIDAD AND TOBAGO: 5 females, OSUC 546023, 546097– 546098, 546101–546102 (CNCI). VENEZUELA: 163 females, 1 male, OSUC 545835–545846, 545853, 545858–545861, 545873–545878, 545902, 545905–545934, 545938, 545944–545947, 545951–545954, 545960–545967, 545979–545990, 557647, 557650, 557661–557663, 557709– 557710 (CNCI); OSUC 146712, 146714, 146716, 146747, 146764, 146768, 146777–146778, 146786, 334303–334308, 334310–334311, 334422–334423, 334511, 334514–334515, 334520– 334522, 334524, 334536, 334539, 334542–334544, 334546, 334549–334551, 45296, 45300, 46280, 46286, 46299, 46532, 46537, 46563, 46580, 48128, 48139, 48146, 48166, 48176, 48315, 48326, 48371, 48387, 48396, 48408, 48426, 48459, 48480, 48492, 48605, 48623, 48628, 48641, 48723, 48839, 48881, 48885, 63892–63893, 63905, 63908, 63911, 63917, 64582 (OSUC).

Calliscelio flavicaudaChen & Johnson, new species

64 http://zoobank.org/9A4B41F8-DDB0-449E-9E73-F5818AC5293A http://bioguid.osu.edu/xbiod_concepts/363560 Figures 1.18A–1.18F.

Description. Body length of female: 1.95–1.99 mm (n=20). Body length of male: 1.90–2.05 mm (n=2). Color of head: brown throughout; orange throughout; orange to pale brown. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: present. Setation of upper frons: with dense, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate. Sculpture of frons below median ocellus: smooth; smooth to coriaceous. Sculpture of posterior vertex: granulate to rugulose. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: largely smooth with small granulate area. Ocular setae: absent. A4 in female: distinctly longer than A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: distinctly longer than wide. Form of male antennal flagellomeres: filiform, A11 approximately 3.0x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: orange throughout; orange to pale brown; dark brown. Color of mesosoma in male: dark brown throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: coriaceous. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: smooth with a longitudinal, median carina. Sculpture of metascutellum in male: smooth. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral to median keel. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: hind femur brown, otherwise yellow. Sculpture of hind coxa: smooth.

65 Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly longer than r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: anterior margin of T2 and T6 yellow, otherwise brown. Color of metasoma in male: anterior margin of T2 yellow, otherwise brown to black. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: transversely striate. Sculpture of posterior margin of T1 in female: smooth medially, longitudinally striate laterally. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially. Sculpture of T3: smooth. Shape of T6 in female: distinctly elongate, approximately 2.0x longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. Females of this species are most similar to C. migma in size and habitus but can be distinguished by the strongly foveolate scutoscutellar sulcus and the smooth T3. Males of C. flavicauda are similar to C. laticintus and C. longius. It may be separated from C. laticinctus by the smooth T3 and the brown hind femora, from C. longiusthat A11 is approximately 3.0x longer than wide. Etymology. The specific epithet meaning "yellow tail" refers to the yellow T6 in female and should be treated as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=363560] Material Examined. Holotype, female: ECUADOR: Sucumbíos Prov., 270m, 00°30'S 76°30'W, Sacha Lodge, 3.VII–13.VII.1994, Malaise trap, P. Hibbs, OSUC 553509 (deposited in CNCI). Paratypes: (52 females, 2 males) BRAZIL: 1 female, OSUC 534533 (CNCI). COLOMBIA: 11 females, OSUC 557602 (CNCI); OSUC 143969, 152156–152157, 178097, 178184, 182734, 189289, 193281, 193905, 262951 (OSUC). ECUADOR: 22 females, 2 males, OSUC 458499, 458509, 458511, 458535, 534224, 534232, 534234, 553377, 553442, 553506– 553508, 553512, 553515, 553517, 553520–553523, 553562, 553566–553567, 553571, 553686 (CNCI). FRENCH GUIANA: 2 females, OSUC 546103 (CNCI); OSUC 570550 (OSUC). PERU: 16 females, OSUC 534417, 534421, 553992, 553994, 554007, 554036, 554038–554042, 554044–554046, 554050 (CNCI); OSUC 343060 (USNM).

Calliscelio foveolatusChen & Johnson, new species

66 http://zoobank.org/5734D0C9-1954-4A66-8E9E-412371890123 http://bioguid.osu.edu/xbiod_concepts/384809 Figures 1.19A–1.19F.

Description. Body length of female: 1.92–2.30 mm (n=10). Color of head: black throughout; reddish orange throughout. Color of antennal clava (A7–A12): black; A7 orange, remainder dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth with sparse punctures. Sculpture of frons below median ocellus: largely smooth with sparse fine punctures. Sculpture of posterior vertex: coriaceous; largely smooth with irregular fine sculpture. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter; greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: slightly longer than wide. Color of mesosoma in female: orange throughout; black throughout; variably yellow to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: coriaceous; coriaceous anteriorly, smooth with sparse fine punctures posteriorly. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: smooth with a longitudinal, median carina. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: greater than 3.0x length of r-rs.

67 Color of metasoma in female: dark brown. Horn on T1 in female: weakly developed. Sculpture of T1 horn dorsally: smooth. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially. Sculpture of T3: smooth with longitudinal submedian striae. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. This species is most similar to C. levis but can be distinguished by the foveolae of the scutoscutellar sulcus between the notauli which are as large as those along the margin of axilla and that A4 is distinctly shorter than A3. Etymology. The epithet is an adjective, derived from the Latin word for small hole, in reference to the sculpture of the scutoscutellar sulcus. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384809] Material Examined. Holotype, female: COSTA RICA: Puntarenas Prov., 24km W Piedras Blancas, Golfo Dulce, 200m, VI.1989–VIII.1989, Hanson & Goulet, OSUC 532869 (deposited in CNCI). Paratypes: (9 females) BOLIVIA: 1 female, OSUC 534177 (CNCI). COSTA RICA: 2 females, OSUC 532802, 532846 (CNCI). ECUADOR: 1 female, OSUC 553581 (CNCI). PERU: 4 females, OSUC 534393–534394, 534402, 553989 (CNCI). VENEZUELA: 1 female, OSUC 545903 (CNCI).

Calliscelio gatineauChen & Johnson, new species http://zoobank.org/0C15B75D-0968-4247-BD87-9C08687543C8 http://bioguid.osu.edu/xbiod_concepts/362062 Figures 1.2A, 1.20A–1.20F.

Description. Body length of female: 2.97 mm (n=1). Color of head: orange throughout. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, short setae. IOS/EH: IOS slightly greater than EH. Sculpture of ventrolateral frons: smooth with sparse punctures. Sculpture of frons below median ocellus: granulate to finely punctate. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate. Ocular

68 setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Color of mesosoma in female: orange throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: rugulose throughout. Sculpture of netrion: rugose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: absent. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior somewhat rounded, approximately 2.5x wider than long. Sculpture of metascutellum in female: rugose. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: orange throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: short, decumbent, hardly exceeding the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as r-rs. Color of metasoma in female: orange throughout. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: granulate medially, with V-shaped keels laterally. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: longitudinally striate throughout. Shape of T6 in female: short, wider than long. Sculpture of S3: longitudinally striate. Diagnosis. This species is most similar to the orange females of C. rubriclavus but can be distinguished based on the smooth sculpture of the mesepisternum below the mesopleural depression and the medially granulate T1 horn , with V-shaped keels laterally. Etymology. The specific epithet refers to the locality on the label of the holotype and should be treated as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362062] Material Examined. Holotype, female: CANADA: QC, path/ridge/woodpile, Gatineau Park, IX–XI.1995, L. Masner, OSUC 534340 (deposited in CNCI).

69 Calliscelio glaberChen & Masner, new species http://zoobank.org/E7A2A6C2-E787-4F5A-BC99-958CF2FF06DF http://bioguid.osu.edu/xbiod_concepts/362055 Figures 1.21A–1.21F.

Description. Body length of female: 1.79–2.17 mm (n=20). Body length of male: 1.80–2.36mm (n=20). Color of head: brown throughout; orange throughout; orange to pale brown. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: glabrous. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth. Sculpture of frons below median ocellus: smooth. Sculpture of posterior vertex: smooth. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: as long as wide. Form of male antennal flagellomeres: filiform, A11 approximately 3.0x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: orange throughout; orange to pale brown. Color of mesosoma in male: orange throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth anteriorly, punctate rugulose posteriorly. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: smooth throughout. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: absent. Sculpture of mesoscutellum: smooth throughout. Shape of metascutellum: posterior margin rounded, approximately 2.5x wider than long. Sculpture of metascutellum in female: smooth. Sculpture of metascutellum in male: smooth. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: orange yellow. Sculpture of hind coxa: smooth.

70 Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: orange throughout. Color of metasoma in male: orange throughout. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: concentrically striate. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: largely smooth with submedian longitudinal striae. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. This species is most similar to C. paraglaber in smooth body sculpture and size but can be distinguished by the interrupted occipital carina (occipital carina complete in C. paraglaber). Etymology. The epithet is an adjective and Latin word for smooth, in reference to the smooth body surface. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362055] Material Examined. Holotype, female: DOMINICAN REPUBLIC: Pedernales Prov., km25 of Alcoa Road, dry montane forest, Baoruco (Bahoruco) Mountains, 700m, 18.I.1989, L. Masner, OSUC 458136 (deposited in CNCI). Paratypes: (27 females, 20 males) CUBA: 2 females, 1 male, OSUC 458145–458146, 458241 (CNCI). DOMINICAN REPUBLIC: 25 females, 19 males, OSUC 458137–458144, 458147–458176, 534372–534373, 534376, 534594– 534596 (CNCI).

Calliscelio granulatusChen & Masner, new species http://zoobank.org/DC45F6BB-FDE6-4E77-8A00-9B2FD32359E4 http://bioguid.osu.edu/xbiod_concepts/362058 Figures 1.22A–1.22F.

Description. Body length of female: 2.31–2.53 mm (n=8). Body length of male: 2.24–2.26mm (n=4). Color of head: orange throughout. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with

71 dense, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate. Ocular setae: dense, short. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Form of male antennal flagellomeres: filiform, A11 approximately 3.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: orange throughout. Color of mesosoma in male: orange throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: largely smooth, granulate ventrally and posteriorly. Sculpture of netrion: rugulose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: orange yellow. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, weakly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: orange with pale brown patch on T1, T2 and T6. Color of metasoma in male: orange with pale brown patch on T2 and T5–T7. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: longitudinally striate. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially; reaching posterior margin of T2. Sculpture of T3: smooth. Shape of T6 in female: distinctly elongate, approximately 2.0x longer than wide. Sculpture of S3: smooth.

72 Diagnosis. This species is most similar to C. telum in hairy compound eyes and habitus but can be distinguished by its rugulose netrion and longitudinally striate T1 horn. Etymology. The adjectival specific epithet refers to the sculpture of the body. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362058] Material Examined. Holotype, female: CUBA: Santiago de Cuba Prov., botanical garden/disturbed and scrub forest, Santiago de Cuba, 5–50m, 5.XII–17.XII.1995, flight intercept trap, S. Peck, OSUC 534599 (deposited in CNCI). Paratypes: CUBA: 7 females, 4 males, OSUC 458304–458314 (CNCI).

Calliscelio laticinctus Ashmead http://zoobank.org/A4202C74-7737-441D-86A1-876224167CFA http://bioguid.osu.edu/xbiod_concepts/4152 Figures 1.2C, 1.23A–1.23F.

Calliscelio laticinctus Ashmead, 1893: 219 (original description); Ashmead, 1894: 223 (redescribed as new); Ashmead, 1900: 327 (distribution); Kieffer, 1926: 499 (description, keyed); Masner, 1976: 38 (description, type information).

Description. Body length of female: 1.97–3.01 mm (n=20). Body length of male: 1.82–2.34mm (n=20). Color of head: yellow throughout. Color of antennal clava (A7–A12): black. Shape of head: subglobose. Central keel of frons: present. Setation of upper frons: with dense, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate. Sculpture of frons below median ocellus: coriaceous. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: as long as A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: distinctly longer than wide. Form of male antennal flagellomeres: filiform, A11 approximately 3.0x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: yellow throughout. Color of mesosoma in male: orange throughout; variably orange to pale brown; brown throughout. Sculpture of dorsal pronotal area:

73 rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: orange throughout; pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, weakly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly longer than r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: dark brown; orange throughout; variably orange to black. Color of metasoma in male: brown throughout; variably yellow to pale brown; black throughout. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: concentrically striate. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: present on anterior margin of T2 medially, reaching posterior margin of T2 laterally. Sculpture of T3: largely smooth with submedian longitudinal striae. Shape of T6 in female: distinctly elongate, approximately 2.5x longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. Females of this species are most similar to C. longius and C. prolixus in color, size and habitus. It can be separated from C. longius by A4 is slightly longer than A3 and A5 shorter than A3, from C. prolixus by the presence of T1 horn and the elongate T6. Males of C. laticinctus are similar to C. flavicauda and C. longius. It may be separated from C. flavicauda by T3 with submedian longitudinal striae and the brown hind femora, from C. longiusby A11 is approximately 3.0x longer than wide. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=4152]

74 Material Examined. Syntype, female, C. laticinctus: SAINT VINCENT AND THE GRENADINES: Saint Vincent Island, no date, H. H. Smith, USNMENT01059105 (deposited in USNM). Syntype, female, C. laticinctus: SAINT VINCENT AND THE GRENADINES: Saint Vincent Island, no date, H. H. Smith, USNMENT01059354 (deposited in USNM). Other material: (323 females, 44 males) BRAZIL: 162 females, 32 males, OSUC 534536, 534538 (CNCI); OSUC 252092, 252094, 252097, 252105–252106, 252137, 252139, 252145, 254591, 322151, 322535–322537, 322539, 322548, 322557, 322720, 323005, 323007, 323080–323081, 323083–323084, 323086–323090, 323092–323093, 323959, 326149, 326194, 326220, 326222, 326235, 343696, 344403, 344481, 345098, 345119, 345205, 345207, 345333, 346169, 346870, 347431, 347438, 347452, 347482, 348267, 348488, 348496, 348669, 348677, 348992, 349017, 349024, 349191, 349194, 349196, 349198, 349513, 349780, 349782, 351241, 352056, 352061– 352062, 352066–352067, 352071, 352249, 352359, 352799, 353038–353039, 353176, 354567, 354919, 355162, 355324–355328, 357042, 361705, 363838, 363840–363842, 363845, 363847, 366692, 366711, 366780, 366782, 366807, 366943, 368420, 370883–370886, 370888–370889, 370891, 370893–370896, 370899, 370906, 370929, 373762, 373770, 374565, 374567, 374575, 374586–374587, 374592, 374597, 374623–374630, 374737, 374742, 375308, 375310, 375315, 376518, 376524, 376976, 376978–376979, 376981–376982, 376985–376988, 377482–377491, 377494, 377504, 378011–378013, 378052, 378065, 378069, 378975–378976, 380179, 380183– 380185, 380187–380189, 380193, 477165, 534688–534691, 55934, 55941, 55943, 577134, 577156, 577159, 577161, 583248–583249 (OSUC); OSUC 225355, 237645 (USNM). COLOMBIA: 39 females, 1 male, OSUC 534554, 557600–557601, 557641 (CNCI); OSUC 152155, 162597, 178015, 178205, 182484, 188778, 189200, 189211, 190987, 190990–190991, 191141, 191180, 191251, 192205, 192209, 193353, 193462–193463, 193547, 232296, 232303, 259575–259576, 259757, 262613, 267804, 269431, 272086, 274573, 274973, 276235, 279926, 279929, 363586, 364067 (OSUC). DOMINICA: 5 females, 1 male, OSUC 458242–458243, 458245–458248 (CNCI). ECUADOR: 4 females, OSUC 458237, 553441, 553477, 557140 (CNCI). FRENCH GUIANA: 31 females, 1 male, OSUC 458388, 458400–458401, 458412, 458434–458435, 458437, 458441–458443, 458449–458453, 458455, 458461, 458463, 458465– 458468, 458470, 458475–458477, 458479, 546105, 546111, 546139, 546150 (CNCI); OSUC 570551 (OSUC). GRENADA: 2 females, OSUC 534253–534254 (CNCI). GUYANA: 11 females, OSUC 458236, 458252–458261 (CNCI). PANAMA: 2 females, OSUC 553757,

75 553771 (CNCI). PERU: 2 females, OSUC 534422, 534426 (CNCI). SAINT VINCENT AND THE GRENADINES: 1 female, OSUC 458244 (CNCI). SURINAME: 2 females, OSUC 534584, 534586 (CNCI). VENEZUELA: 62 females, 9 males, OSUC 534314–534339, 545956, 557659–557660, 557667, 557681, 557683, 557685–557695, 557698 (CNCI); OSUC 146713, 146715, 146741, 146760, 334309, 334312, 334344, 334347–334348, 334438, 334513, 334516, 334527, 334538, 334552, 360647, 360650, 360738, 367061–367062, 48118, 48901, 576990– 576992, 64571, 79763 (OSUC).

Calliscelio latifronsChen & Johnson, new species http://zoobank.org/F45AC467-663F-4811-B40F-AFC2493439BE http://bioguid.osu.edu/xbiod_concepts/367272 Figures 1.1D–1.1F, 1.24A–1.24F.

Description. Body length of female: 1.85–2.50 mm (n=11). Body length of male: 1.70–2.05mm (n=10). Color of head: black throughout. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, short setae. IOS/EH: IOS slightly greater than EH. Sculpture of ventrolateral frons: granulate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate. Ocular setae: sparse, short. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: as long as wide. Form of male antennal flagellomeres: moniliform, A11 as long as wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: black throughout; orange to pale brown. Color of mesosoma in male: dark brown throughout. Sculpture of dorsal pronotal area: granulate. Sculpture of lateral pronotal area: granulate throughout. Sculpture of netrion: rugulose. Notaulus: percurrent; abbreviated, at most reaching middle of mesoscutum. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: transverse. Foveolae of scutoscutellar sulcus between notauli: absent. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: granulate. Sculpture

76 of metascutellum in male: granulate. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: granulate. Sculpture of ventral metapleural area: smooth dorsally, granulate ventrally. Color of legs: orange throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: variably orange to pale brown. Color of metasoma in male: variably orange to pale brown; pale brown throughout. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: granulate. Sculpture of posterior margin of T1 in female: striate rugose. Sculpture of T1 in male: longitudinally rugose. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth medially, longitudinally striate laterally. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: granulate. Diagnosis. This species is most similar to C. armila but can be easily distinguished by the granulate upper frons and the longitudinally striate T3. Etymology. The epithet is a compound noun in apposition referring to the wide IOS. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=367272] Material Examined. Holotype, female: PARAGUAY: Presidente Hayes Dept., 151m, 23°48'S 60°46'W, Escalante Lagoon, 27.XI.2003, yellow pan trap, B. Garcete, OSUC 323077 (deposited in MNHNPY). Paratypes: (10 females, 10 males) ARGENTINA: 2 females, OSUC 534438–534439 (CNCI). BRAZIL: 1 female, OSUC 534517 (CNCI). PARAGUAY: 7 females, 10 males, OSUC 322992, 363637, 534686-534687 (MNHNPY); OSUC 150574-150575, 276703, 322991, 322993, 323075-323076, 323078, 363707, 363711-363712, 434082-434083 (OSUC).

Calliscelio levisChen & Johnson, new species http://zoobank.org/06FA4764-0408-4049-9F2C-A5997337F090

77 http://bioguid.osu.edu/xbiod_concepts/384811 Figures 1.25A–1.25F.

Description. Body length of female: 1.52–1.77 mm (n=13). Color of head: variably brown to black. Color of antennal clava (A7–A12): A7 orange, remainder dark brown to black; A7 dark brown, A8–A12 golden yellow. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth with sparse punctures. Sculpture of frons below median ocellus: smooth. Sculpture of posterior vertex: coriaceous. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: as long as A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: distinctly longer than wide. Color of mesosoma in female: orange throughout; variably yellow to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: rugulose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: coriaceous. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin somewhat rounded, approximately 3.5x wider than long. Sculpture of metascutellum in female: smooth. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange yellow. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: variably yellow to pale brown. Horn on T1 in female: weakly developed. Sculpture of T1 horn dorsally: smooth. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Development of longitudinal striae on T2 in female:

78 present on anterior margin of T2 medially, reaching posterior margin of T2 laterally. Sculpture of T3: smooth; smooth with longitudinal submedian striae. Shape of T6 in female: short, subtriangular. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. foveolatus but can be distinguished because the foveolae of the scutoscutellar sulcus between the notauli are smaller than those along the margin of the axilla and A4 is as long as A3. Etymology. The epithet is an adjective, Latin word for smooth, polished and bald, in reference to the smooth T3. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384811] Material Examined. Holotype, female: ECUADOR: Napo Prov., Baeza, 2000m, 20.II– 25.II.1979, Malaise trap, W. Mason, OSUC 553454 (deposited in CNCI). Paratypes: (12 females) ECUADOR: 10 females, OSUC 458530–458531, 553452, 553455, 553484, 553573, 553579–553580, 553582, 553691 (CNCI). PERU: 2 females, OSUC 534395, 534415 (CNCI).

Calliscelio longiusChen & Johnson, new species http://zoobank.org/36D661EE-96E2-4EB5-BB2F-0056C9FF514B http://bioguid.osu.edu/xbiod_concepts/384815 Figures 1.1J–1.1L, 1.26A–1.26F.

Description. Body length of female: 2.23–2.97 mm (n=20). Body length of male: 1.86–1.91mm (n=3). Color of head: black throughout; orange throughout; variably brown to black; variably orange to dark brown. Color of antennal clava (A7–A12): black. Shape of head: subglobose. Central keel of frons: present. Setation of upper frons: with dense, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate. Sculpture of frons below median ocellus: coriaceous; smooth to coriaceous. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly longer than A3. A5 in female: longer than A3, distinctly longer than wide. Shape of female A6: length distinctly greater than width. Form of male antennal flagellomeres: filiform, A11 approximately 4.5x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5.

79 Color of mesosoma in female: orange throughout; black throughout; variably orange to pale brown. Color of mesosoma in male: brown throughout; black throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: coriaceous; granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: smooth with a longitudinal, median carina. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral to median keel. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: orange yellow; hind femur brown, otherwise yellow. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly longer than r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: dark brown; orange throughout; variably orange to black. Color of metasoma in male: brown throughout; variably yellow to pale brown; black throughout. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: transversely striate; smooth to somewhat transversely striate. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2; present on anterior margin of T2 medially, reaching posterior margin of T2 laterally. Sculpture of T3: smooth; smooth with longitudinal submedian striae. Shape of T6 in female: distinctly elongate, approximately 3.5x longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. The females of this species are most similar to C. laticinctus and C. prolixus in color, size and habitus. It can be separated from C. laticinctus by A4 distinctly longer than A3 and A5 longer than A3, from C. prolixus by the presence of T1 horn and the elongate T6.

80 Etymology. The epithet is an adjective, the Latin word for longer, in reference to A5 longer than A3. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384815] Material Examined. Holotype, female: COLOMBIA: Nariño Dept., Permanent Parcel, M.918, 1885m, 01°15'N 78°15'W, La Planada Nature Reserve, 16.VIII–2.IX.2000, Malaise trap, G. Oliva, OSUC 374730 (deposited in IAVH). Paratypes: (102 females, 3 males) BRAZIL: 3 females, OSUC 534528–534529, 534532 (CNCI). COLOMBIA: 62 females, 1 male, OSUC 557598 (CNCI); OSUC 152153, 152158-152159, 162497, 162588-162589, 162600, 178167, 178187, 178190, 182483, 182735, 182749-182751, 188924, 188926, 188947, 188952, 188956, 188958, 189208, 189212, 189293, 189300, 190935, 191037-191038, 191041, 191301, 191305, 191316, 191318, 193200, 193299, 193319, 193544, 193561-193562, 193567-193569, 193581- 193582, 193596, 193685-193686, 193832, 193841, 193915, 193935, 262614-262615, 274571, 274574, 275802, 276238, 372641, 374729, 374731-374732, 377427 (OSUC).ECUADOR: 18 females, 1 male, OSUC 458238, 458487, 553371, 553436, 553444–553445, 553463, 553467, 553474, 553476, 553519, 553527, 553538, 553546, 553596, 553606, 553678, 553690, 557141 (CNCI). PERU: 17 females, 1 male, OSUC 534406, 534413, 534423–534425, 553964, 554034, 554037, 554048, 554051, 554054 (CNCI); OSUC 323931 (OSUC); OSUC 228066–228067, 228070, 228134, 231999, 343061 (USNM). VENEZUELA: 2 females, OSUC 557699–557700 (CNCI).

Calliscelio magnificus Chen & Masner, new species http://zoobank.org/809ACB6E-A75D-4C0F-8F0E-1643F54AE167 http://bioguid.osu.edu/xbiod_concepts/362060 Figures 1.27A–127F.

Description. Body length of female: 2.85–3.88 mm (n=19). Body length of male: 2.86–3.00mm (n=7). Color of head: dark brown; orange to pale brown. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: present. Setation of upper frons: with sparse, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth to rugulose. Sculpture of frons below median ocellus: smooth; coriaceous. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially:

81 complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: largely smooth with small granulate area. Ocular setae: absent. A4 in female: distinctly longer than A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: length distinctly greater than width. Form of male antennal flagellomeres: filiform, A11 approximately 3.5x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: variably orange to pale brown. Color of mesosoma in male: variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: broad, short. Sculpture of metascutellum in female: smooth with a longitudinal, median carina. Sculpture of metascutellum in male: smooth. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral median keel. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: orange yellow; pale brown. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: variably orange to pale brown. Color of metasoma in male: variably orange to pale brown. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: smooth. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: present on anterior margin of T2 medially, reaching posterior margin of T2 laterally. Sculpture of T3: smooth with longitudinal submedian striae. Shape of T6 in female: distinctly elongate, approximately 3.0x longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures.

82 Diagnosis. The females of this species are easily distinguished from other Calliscelio species by the combination of large and smooth T1 horn, elongate T6 and large body size. The males are most similar to C. flavocauda, C. laticinctus and C. longius but can be separated by the smooth metascutellum. Etymology. The specific epithet is to be treated as a noun in apposition, derived from the Latin for "splendid" and refers to our impression of this species. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362060] Material Examined. Holotype, female: PERU: Huánuco Reg., 39km NE Tingo Maria, Azul Range, 1700m, 11.I–14.I.1983, Newton & Thayer, OSUC 458272 (deposited in CNCI). Paratypes: (20 females, 8 males) COSTA RICA: 2 females, OSUC 532663, 532746 (CNCI). ECUADOR: 1 female, OSUC 553449 (CNCI). PERU: 13 females, 8 males, OSUC 458263– 458264, 458266–458271, 458273–458284, 534405 (CNCI). VENEZUELA: 4 females, OSUC 458233–458235, 557668 (CNCI).

Calliscelio migmaChen & Johnson, new species http://zoobank.org/2F3A637F-06EC-4ECE-8E73-D337BA903320 http://bioguid.osu.edu/xbiod_concepts/384750 Figures 1.28A–1.28F.

Description. Body length of female: 1.34–2.16 mm (n=20). Body length of male: 1.43–1.94mm (n=20). Color of head: yellow throughout; orange throughout. Color of antennal clava (A7–A12): dark brown to black; A7, A8 brown, A9–A12 white to pale yellow. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with dense, short setae; with sparse, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth to granulate. Sculpture of frons below median ocellus: smooth; granulate. Sculpture of posterior vertex: smooth; granulate to rugulose. Hyperoccipital carina: absent. Occipital carina medially: weakly indicated, irregularly sculptured. Length of OOL: less than 0.5x ocellar diameter; greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly longer than A3. Shape of female A5: length distinctly greater than width. Shape of female A6: length distinctly greater than width. Form of male antennal flagellomeres:

83 thread-like, length at least 4.0x greater than width. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: orange throughout; yellow throughout. Color of mesosoma in male: orange throughout; yellow throughout. Sculpture of dorsal pronotal area: rugose; smooth. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: smooth with sparse punctures; coriaceous. Shape of mesoscutellum: semiellipsoidal. Scutoscutellar sulcus medially: weakly foveolate. Sculpture of mesoscutellum: smooth with sparse fine punctures; coriaceous. Shape of metascutellum: broad, short. Sculpture of metascutellum in female: smooth; rugose. Sculpture of metascutellum in male: rugose; smooth. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of lateral propodeal area in female: rugose. Sculpture of lateral propodeal area in male: rugose with one or two longitudinal keels lateral to median keel; longitudinally striate. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth; largely smooth, rugose ventrally. Color of legs: orange throughout; pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: orange throughout; yellow throughout. Color of metasoma in male: orange throughout; variably orange to pale brown. Horn on T1 in female: large and distinct; present as a small bulge. Sculpture of T1 horn dorsally: smooth; rugulose; transversely striate; smooth to somewhat transversely striate. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: present on anterior margin of T2 medially, reach posterior margin of T2 laterally. Sculpture of T3: smooth with longitudinal submedian striae; longitudinally striate throughout. Shape of T6 in female: short, subtriangular. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. This species is most similar to C. amadoi in color and size but it can be easily distinguished by having A5 and A6 in female distinctly longer than wide, whereas in C. amadoi A5 is slightly longer than wide, A6 quadrate.

84 Etymology. The epithet is used as a noun in apposition derived from the Greek word for mixture, in reference to the variations in female antennae color and T1 horn sculpture. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384750] Material Examined. Holotype, female: BRAZIL: PA, Melgaço Mpio., Igarapé Tijucaquara, trail, M11, P05128, 01°44'12.8"S 51°29'56.6"W, Caxiuanã, 18.XI–24.XI.2003, Malaise trap, A. P. Aguiar & J. Dias, OSUC 380181 (deposited inMPEG). Paratypes: (661 females, 82 males) BOLIVIA: 1 female, OSUC 534172 (CNCI). BRAZIL: 233 females, 41 males, OSUC 534513-534516, 534531, 534535, 534543-534545, 557166, 557201-557202, 557204, 557208-557209, 557395 (CNCI); OSUC 252141, 322371, 322377, 322671, 326226, 326230, 344407-344408, 344487, 345100, 345312, 345325, 345331, 345589, 346174, 346180, 346888, 346946, 346954, 346964, 347060, 347207-347208, 347213, 347229, 347427, 347437, 348008, 348115, 348641, 348678, 349368, 349636, 349659, 349685, 349771, 349783, 349785, 351289, 351329, 351462, 351467, 351525, 351528, 351673, 351677-351678, 351680-351683, 351815, 351823, 351833, 351836-351838, 352357, 352800, 352802, 352806, 352810, 352812, 353174-353175, 353177, 353717, 353725-353726, 353732, 353951-353953, 354075, 354082, 354710, 354831, 354885, 354892, 354895, 354898, 354900, 362588-362589, 363839, 363843- 363844, 363849, 370897, 370905, 370908, 370915, 370923-370924, 371841-371842, 371844, 376057, 376541, 378963, 477166-477167, 577157, 577162, 577173, 583256 (MZSP); OSUC 251768, 252087, 322150, 322152, 322370, 322532, 322545-322547, 323091, 323958, 323964, 323970, 326189, 326397, 326404, 326502-326503, 326505, 326538, 337221-337222, 346041, 346997, 347244, 347277, 347293, 347295, 347655, 347660, 347671, 347678, 347852, 347871, 347954, 348074-348075, 348081, 348094, 348097, 348117, 348356, 349102-349103, 349107, 349111, 349201, 351425, 353047, 353426, 354797, 357021, 362607, 366698, 366701, 366712, 366721, 366778-366779, 366790, 366808, 366938-366939, 366944-366945, 366953, 366964- 366966, 366974, 371836, 371839, 371849, 372549-372550, 373757, 373761, 373773, 373779, 374557, 374560-374561, 374717, 374739-374741, 376486, 376498-376500, 376505, 376509, 376527, 376530-376531, 376538, 376542, 376966, 377495-377497, 378014-378015, 378033, 378035, 378041, 378054-378055, 378063, 378080, 378085, 378595, 378597, 378960-378962, 378967, 378969, 378972-378973, 378979, 380182, 45697, 45721, 55930-55932, 55946, 55953, 61378, 61404, 61446, 61454, 61499, 61504, 61518, 61580, 63283, 63288, 63292, 63295, 63309, 63319, 63330, 63353, 63362, 63431, 63439, 63538, 63643-63644, 63665 (OSUC).

85 COLOMBIA: 330 females, 19 males, OSUC 557547, 557549, 557555-557556, 557599, 557622, 557640, 557645-557646 (CNCI); OSUC 144165, 152145, 152147-152148, 162500, 162506, 162510, 162514, 162590, 162592, 162605, 162614, 162621-162623, 178018-178019, 178091, 178157-178159, 178161, 178165-178166, 178169-178173, 178175-178177, 178183, 178185, 178188-178189, 178194, 178196-178197, 178199-178200, 178202-178204, 178206, 179458-179459, 182585-182587, 182589-182590, 182594, 182721, 182736, 182738-182739, 188624, 188684, 188921, 188923, 188928-188930, 188950, 188953, 189276, 189278, 189280, 189284, 189288, 189301-189302, 189392, 191095, 193318, 193323, 193332, 193335-193337, 193339, 193341, 193343, 193345, 193354, 193356, 193359, 193394, 193396-193397, 193400, 193403, 193537, 193540, 193546, 193590, 231806-231809, 231813-231817, 231819, 231822, 231828, 232300, 259764, 272088, 275806-275807, 275809, 276187, 276195-276196, 276240, 278953, 279660, 363589, 363599, 364074, 372635 (IAVH); OSUC 143967-143968, 143972, 152133-152140, 152142-152144, 152146, 162498, 162502, 162585, 162595, 162598, 162604, 176896, 178164, 182225-182226, 182488-182490, 182498, 182500, 182593, 182744-182748, 182763-182765, 182767, 188549, 188681, 188687, 188922, 188925, 188927, 188931-188932, 188935-188936, 188938-188939, 189177-189178, 189180-189182, 189184, 189187-189189, 189192, 189194, 189196-189197, 189201-189202, 189204, 189206-189207, 189209, 189214, 189281, 189286, 189298-189299, 190307-190309, 190312, 191039, 191177-191179, 191181, 191186, 191207, 191319, 191826, 193277, 193279-193280, 193283, 193285-193287, 193290, 193333-193334, 193344, 193346, 193350-193352, 193357, 193398-193399, 193402, 193405- 193406, 193412, 193423, 193468-193470, 193539, 193545, 193550, 193571, 193573-193574, 193577-193580, 193585-193586, 193588, 193592, 193594, 193777, 193780, 193901, 193904, 193907-193909, 193911-193912, 193914, 193934, 193936-193937, 193939, 194198, 231805, 231823, 231825-231826, 249898, 253452, 253456, 253461, 259751, 259755, 259763, 262602- 262605, 262617-262618, 262942-262944, 262946-262947, 262949-262950, 262952, 262955- 262960, 262965, 267810, 269349, 269351, 269354, 269433-269435, 269437-269439, 269483, 272082, 272084, 272091, 273456, 274971, 275028-275029, 275812-275815, 275857-275859, 276192, 276233-276234, 279656, 279659, 363588, 364078, 372632-372633, 372637-372638, 372643, 76996 (OSUC). COSTA RICA: 1 male, OSUC 245173 (OSUC). ECUADOR: 41 females, 3 males, OSUC 458485, 458491–458493, 532693–532694, 534223, 534230–534231, 534233, 534243, 534246–534247, 553247, 553385, 553390–553391, 553393–553395, 553397,

86 553405, 553415, 553457–553460, 553549, 553557, 553563–553564, 553585, 553592, 553627– 553628, 553682, 553730, 577332 (CNCI); OSUC 534657, 534664–534667, 534671 (OSUC). FRENCH GUIANA: 26 females, 10 males, OSUC 458389, 458391, 458399, 458407–458408, 458417–458420, 458424–458425, 458430–458433, 458436, 458438–458440, 458447–458448, 458454, 458456–458458, 533981, 534553, 546110, 546129, 546131, 546133–546134, 546142– 546143, 546146–546147 (CNCI). PERU: 30 females, 8 males, OSUC 534390, 534400, 534414, 534418, 534420, 534429, 553954, 553957–553961, 553963, 553966, 553978, 553990, 554000, 554002, 554005–554006, 554008–554009, 554030, 554033, 554053 (CNCI); OSUC 323932, 570535–570536, 570545, 570547–570548 (OSUC); OSUC 228068–228069, 228071, 228074, 228193–228194, 232000 (USNM). Comments. This species is well supported by many characters, although there are variations in female antennae color and T1 horn sculpture. The common color of the female antennal club is dark brown to black, while there are a few exceptions that A7, A8 brown, A9– A12 white to pale yellow. T1 horn usually is smooth, while a few species rugulose, or transversely striate, or smooth to somewhat transversely striate. These variations are gradual among specimens. Therefore we consider them as intraspecific rather than interspecific.

Calliscelio minutiaChen & Johnson, new species http://zoobank.org/89839644-E7FF-40CF-834B-5C380C230B9B http://bioguid.osu.edu/xbiod_concepts/363279 Figures 1.29A–1.29F.

Description. Body length of female: 1.70–2.06 mm (n=20). Body length of male: 1.77–1.94mm (n=8). Color of head: yellow throughout. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Form of male antennal flagellomeres: filiform,

87 A11 approximately 2.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: yellow throughout; yellow with mesoscutellum pale brown. Color of mesosoma in male: yellow throughout; variably yellow to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth anteriorly, granulate posteriorly. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: absent. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin rounded, approximately 3.0x wider than long. Sculpture of metascutellum in female: smooth. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: smooth to rugulose. Sculpture of dorsal propodeum in male: rugulose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth dorsally, densely punctate ventrally. Color of legs: pale yellow throughout; hind coxa pale brown, otherwise pale yellow. Sculpture of hind coxa: smooth. Color of fore wing: hyaline with infuscate band in the middle. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: yellow with variable pale brown patches; yellow throughout. Color of metasoma in male: variably yellow to pale brown. Horn on T1 in female: absent. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth. Shape of T6 in female: short, wider than long. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. suni in color, size and habitus, and it is also similar to small specimens of C. sfina. It can be separated by the presence of an infuscate band in the middle of the fore wing. Etymology. The epithet is used as a noun in apposition derived from the Latin word for smallness, in reference to the body size. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=363279]

88 Material Examined. Holotype, female: BRAZIL: BA, Mata da Esperança, YPT T3, 14°46'S 39°04'W, Ilheus, 18.V–21.V.2002, yellow pan trap, A. M. Penteado-Dias et al., OSUC 433847 (deposited in MZSP). Paratypes: (23 females, 8 males) BRAZIL: 23 females, 7 males, OSUC 534541 (CNCI); OSUC 433803, 433818 (MZSP); OSUC 150797–150798, 150800, 150965, 150973, 322554–322556, 367440–367441, 378985, 427459–427460, 427462–427463, 433806–433807, 433820, 433822, 433826–433827, 433849–433851, 577073, 583209, 583246 (OSUC). COLOMBIA: 1 male, OSUC 193408 (OSUC).

Calliscelio paraglaberChen & Johnson, new species http://zoobank.org/4C30C899-9BA9-429D-A1EB-C1D4DA175750 http://bioguid.osu.edu/xbiod_concepts/362057 Figures 1.30A–1.30F.

Description. Body length of female: 1.73–2.27 mm (n=20). Body length of male: 1.60–1.63 mm (n=14). Color of head: black throughout. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth. Sculpture of frons below median ocellus: smooth. Sculpture of posterior vertex: smooth. Hyperoccipital carina: absent. Occipital carina medially: weakly indicated, irregularly sculptured. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: as long as wide. Form of male antennal flagellomeres: filiform, A11 approximately 4.0x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: black throughout. Color of mesosoma in male: black throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: smooth throughout. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: smooth throughout. Shape of metascutellum: posterior margin somewhat rounded, approximately 4.0x wider than long. Sculpture of metascutellum in female: smooth.

89 Sculpture of metascutellum in male: smooth. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, weakly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: T1 yellow, otherwise dark brown to black. Color of metasoma in male: brown throughout. Horn on T1 in female: weakly developed. Sculpture of T1 horn dorsally: smooth. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially; reaching posterior margin of T2. Sculpture of T3: smooth with longitudinal submedian striae. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. This species is most similar to C. glaber and C. torqueo in its smooth head and metascutellum. It may be separated from those two species by the complete occipital carina. Etymology. The name refers to the high degree of similarity with C. glaber and is intended to be used as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362057] Material Examined. Holotype, female: COSTA RICA: Puntarenas Prov., 24km W Piedras Blancas, Golfo Dulce, 200m, VI.1989–VIII.1989, Hanson & Goulet, OSUC 532684 (deposited in CNCI). Paratypes: (41 females, 14 males) COLOMBIA: 3 females, OSUC 191100–191101, 269212 (OSUC). COSTA RICA: 32 females, 14 males, OSUC 532459– 532460, 532462, 532491, 532507–532510, 532517, 532521, 532527–532530, 532533, 532551, 532553–532554, 532576, 532589, 532592–532594, 532597–532601, 532610, 532612–532613, 532667–532669, 532673, 532677, 532682, 532686, 532688–532689, 532691–532692, 532708, 532783, 532791, 532929 (CNCI). GUYANA: 2 females, OSUC 534274–534275 (CNCI). MEXICO: 2 females, OSUC 534013, 557255 (CNCI). VENEZUELA: 2 females, OSUC 557665, 557671 (CNCI).

90

Calliscelio pararemigioChen & Masner, new species http://zoobank.org/D91754CD-B065-431B-B920-3428604B4DF5 http://bioguid.osu.edu/xbiod_concepts/362061 Figures 1.31A–1.31F.

Description. Body length of female: 2.53–3.38 mm (n=10). Body length of male: 2.11–2.75mm (n=5). Color of head: black throughout. Color of antennal clava (A7–A12): dark brown to black; A12 pale yellow, remainder dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth with sparse punctures. Sculpture of frons below median ocellus: smooth. Sculpture of posterior vertex: smooth. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: as long as A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: distinctly longer than wide. Form of male antennal flagellomeres: filiform, A11 approximately 3.0x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: orange throughout; variably yellow to pale brown. Color of mesosoma in male: orange throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth anteriorly, granulate posteriorly. Sculpture of netrion: rugulose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: smooth with sparse punctures; coriaceous; densely punctate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin somewhat rounded, approximately 4.0x wider than long. Sculpture of metascutellum in female: finely crenulate. Sculpture of metascutellum in male: finely crenulate. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral median keel. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area:

91 smooth. Color of legs: coxae to femur white, remainder of the legs pale yellow; pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: orange throughout; variably orange to pale brown. Color of metasoma in male: orange throughout; variably orange to pale brown. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: densely and concentrically striate anteriorly, smooth posteriorly. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth with longitudinal submedian striae. Shape of T6 in female: distinctly elongate, approximately 3.0x longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. This species is most similar to C. remigio but can be distinguished by its subglobose head and densely and concentrically striate T1 horn on anterior portion in female. In males, it can be separated from C. remigio by the subglobose head and the largely smooth S3. Etymology. The specific epithet is a reference to the high degree of similarity with C. remigio and is intended to be used as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362061] Material Examined. Holotype, female: CUBA: Santiago de Cuba Prov., La Isabelica, environs of Gran Piedra Mountain, 1100m, 6.XII–7.XII.1995, screen sweeping, L. Masner, OSUC 458239 (deposited in CNCI). Paratypes: (9 females, 5 males) CUBA: 1 female, OSUC 458240 (CNCI). DOMINICAN REPUBLIC: 8 females, 5 males, CMNH-486,653, 490,811 (CMNH); OSUC 458302, 458315–458324 (CNCI).

Calliscelio prolixusChen & Johnson, new species http://zoobank.org/E287C8E0-BBF5-4850-914A-7B0C4BC1548F http://bioguid.osu.edu/xbiod_concepts/384175 Figures 1.32A–1.32F.

92 Description. Body length of female: 2.00–2.48 mm (n=11). Color of head: dark brown; orange to pale brown. Color of antennal clava (A7–A12): dark brown to black; A7 yellow, remainder dark brown to black. Shape of head: subglobose. Central keel of frons: present. Setation of upper frons: with dense, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate. Sculpture of frons below median ocellus: smooth to coriaceous. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly longer than A3. A5 in female: longer than A3, distinctly longer than wide. Shape of female A6: distinctly longer than wide. Color of mesosoma in female: variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: coriaceous. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: rugose. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose with one or two longitudinal keels lateral median keel. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: variably orange to pale brown. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, strongly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly longer than r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: variably yellow to pale brown. Horn on T1 in female: absent. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially. Sculpture of T3: smooth. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: smooth.

93 Diagnosis. This species is most similar to C. longius in elongate antenna, color and habitus. It can be distinguished by the absence of T1 horn and the subtriangular T6 in female. Etymology. The epithet is an adjective, Latin word for long, in reference to the elongate antenna. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384175] Material Examined. Holotype, female: COLOMBIA: Valle del Cauca Dept., 650m, 03°26'N 76°48'W, Farallones de Cali Natural National Park, 19.XII–2.I.2001, Malaise trap, S. Sarria, OSUC 557587 (deposited in CNCI). Paratypes: (10 females) BRAZIL: 1 female, OSUC 322538 (OSUC). COLOMBIA: 4 females, OSUC 557588–557589, 557591 (CNCI); OSUC 268908 (OSUC). ECUADOR: 5 females, OSUC 553448, 553453, 553472, 553516, 553656 (CNCI).

Calliscelio punctatifronsChen & Johnson, new species http://zoobank.org/ACEE562D-FAE5-4C8C-8DE7-92FE3B7ADF08 http://bioguid.osu.edu/xbiod_concepts/363280 Figures 1.1A–1.1C, 1.33A–1.33F.

Description. Body length of female: 1.80–2.56 mm (n=20). Body length of male: 1.45–1.88mm (n=20). Color of head: black throughout. Color of antennal clava (A7–A12): black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with dense, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: irregularly punctate. Sculpture of frons below median ocellus: densely punctate. Sculpture of posterior vertex: densely punctate. Hyperoccipital carina: absent. Occipital carina medially: complete, strongly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate with sparse punctures. Ocular setae: dense, long. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wide than long. Form of male antennal flagellomeres: filiform, A11 approximately 2.5x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: black throughout. Color of mesosoma in male: black throughout. Sculpture of dorsal pronotal area: areolate. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: striate. Notaulus: percurrent or nearly so. Sculpture of

94 mesoscutum: smooth with sparse punctures; coriaceous anteriorly, smooth with sparse fine punctures posteriorly. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 2.5x wider than long. Sculpture of metascutellum in female: rugose; rugose anteriorly, smooth posteriorly. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: largely smooth with a row of foveae along mesopleural carina. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange throughout; pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: subhyaline. Rs+M: nebulose, weakly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: orange throughout; reddish brown to black. Color of metasoma in male: variably orange to pale brown; brown throughout; reddish brown. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: smooth. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially. Sculpture of T3: smooth. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. This species (especially small specimens) is most similar to C. bisulcatus but can be distinguished by the long setae on upper frons and pigmented Rs+M. Etymology. The specific epithet refers to the punctate frons and should be treated as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=363280] Material Examined. Holotype, female: COLOMBIA: Bolívar Dept., La Suiris, M.617, 126m, 09°54'N 75°07'W, Los Colorados Fauna and Flora Sanctuary, 12.IX–15.IX.2000, Malaise trap, E. Deulufeut, OSUC 191124 (deposited in IAVH). Paratypes: (192 females, 55 males) BELIZE: 32 females, 13 males, OSUC 534198, 534201, 534204–534205, 534211–534213,

95 534219–534221, 534276, 534278, 534280–534282, 534285–534289, 534293–534300, 534309, 534312 (CNCI); OSUC 185848, 185852, 243690–243691, 48053, 91681, 91684, 91688, 91691, 91697–91698, 91701, 91704, 93637–93638 (OSUC). BOLIVIA: 4 females, OSUC 534185, 534187, 534190–534191 (CNCI). BRAZIL: 1 female, 7 males, OSUC 557160–557165, 557296 (CNCI); OSUC 133029 (OSUC). COLOMBIA: 20 females, 14 males, OSUC 557409-557413, 557422-557427, 557458-557470, 557609, 557611, 557619 (CNCI); OSUC 191128, 191140 (IAVH); OSUC 182753, 191123, 191125, 191142, 259758 (OSUC).COSTA RICA: 37 females, OSUC 532465, 532469–532470, 532474, 532487–532490, 532497, 532502, 532543, 532558– 532560, 532563, 532580–532582, 532623, 532625, 532640–532641, 532644, 532647–532650, 532653, 532697, 532775, 532781, 532823–532824, 532924, 532930, 534137 (CNCI); OSUC 237331 (OSUC). ECUADOR: 49 females, 4 males, OSUC 458482, 458484, 458495, 458537– 458542, 534238, 553372, 553381, 553413, 553416, 553427–553430, 553432, 553479–553482, 553494, 553528, 553545, 553605, 553630, 553652, 553657–553666, 553668, 553670–553671, 553677, 553689, 553701–553704, 553718, 553731–553732, 553736, 553738 (CNCI). HONDURAS: 11 females, OSUC 399378, 399384–399385, 399392, 399394, 399396, 399400, 399405, 410439, 410442, 410446 (MZLU). MEXICO: 19 females, 5 males, OSUC 532626, 534015–534017, 534021–534023, 534026, 534461–534462, 534471, 534476, 534483–534486 (CNCI); OSUC 375848–375852, 415020, 55929, 55951 (OSUC). PANAMA: 3 females, 10 males, OSUC 534065, 553822, 553892–553894, 553896–553899, 553910, 553912, 553922, 553948 (CNCI). PERU: 6 females, 2 males, OSUC 534396–534397, 534399, 534401, 553968, 553988, 554026, 554028 (CNCI). TRINIDAD AND TOBAGO: 8 females, OSUC 545997, 546030–546032, 546084, 557319–557321 (CNCI). URUGUAY: 2 females, OSUC 534608– 534609 (CNCI).

Calliscelio remigioChen & Masner, new species http://zoobank.org/CB62A4C2-F171-4C23-B065-A63A0A197B7A http://bioguid.osu.edu/xbiod_concepts/362059 Figures 1.34A–1.34F.

Description. Body length of female: 2.56–2.96 mm (n=16). Body length of male: 2.45–2.70mm (n=7). Color of head: black throughout. Color of antennal clava (A7–A12): dark brown to black.

96 Shape of head: strongly transverse. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth with sparse punctures. Sculpture of frons below median ocellus: smooth. Sculpture of posterior vertex: smooth. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: as long as A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: distinctly longet than wide. Form of male antennal flagellomeres: filiform, A11 approximately 3.0x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: orange with longitudinal, median black strip on anterior mesoscutum. Color of mesosoma in male: orange with longitudinal, median black strip on anterior mesoscutum. Sculpture of dorsal pronotal area: smooth. Sculpture of lateral pronotal area: smooth anteriorly, punctate rugulose posteriorly. Sculpture of netrion: rugulose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: smooth with sparse punctures; densely punctate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin straight, approximately 3.0x wider than long. Sculpture of metascutellum in female: finely crenulate. Sculpture of metascutellum in male: finely crenulate. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral median keel. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: coxae to femur white, remainder of the legs pale yellow; pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: variably orange to pale brown. Color of metasoma in male: orange throughout; variably orange to pale brown. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: transversely striate. Sculpture of posterior margin of T1 in female:

97 longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth with longitudinal submedian striae. Shape of T6 in female: distinctly elongate, approximately 3.0x longer than wide. Sculpture of S3: densely punctate; punctate rugose. Diagnosis. This species is most similar to C. pararemigio but can be distinguished by its strongly transverse head and transversely striate small T1 horn in female. In males, it can be separated from C. pararemigio by the strongly transverse head and the densely punctate or punctate rugose S3. Etymology. The specific epithet refers to the locality, Mt. Remigio, on the label of the holotype and should be treated as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362059] Material Examined. Holotype, female: DOMINICAN REPUBLIC: Barahona Prov., Baoruco (Bahoruco) Mts., cloud forest, DR-13, Remigio Knoll, 800m, 26.III.1991, L. Masner, OSUC 458296 (deposited in CNCI). Paratypes: DOMINICAN REPUBLIC: 15 females, 7 males, CMNH-486,650, 486,686, 486,897 (CMNH); OSUC 458285–458295, 458297–458301, 458303, 534377–534378 (CNCI).

Calliscelio rubriclavus (Ashmead), comb. n. http://zoobank.org/B78820F5-FCE1-4A15-BC59-12B9A1C1AAE4 http://bioguid.osu.edu/xbiod_concepts/4169 Figures 1.35A–1.37F.

Acolus rubriclavus Ashmead, 1887: 99 (original description). Anteris nigriceps Ashmead, 1893: 225, 226 (original description); Muesebeck, 1958: 93 [junior synonym of Ceratoteleia marlatti (Ashmead)], syn. n. http://zoobank.org/9C37051A-432A-4772-A8EC-C1345A2F32D4 http://bioguid.osu.edu/xbiod_concepts/8446 Caloteleia Marlattii Ashmead, 1893: 212, 214 (original description, keyed); Harrington, 1900: 187 (variation); Brues, 1903: 126 (emendation of male description), syn. n. http://zoobank.org/91C406E3-85AA-4C93-8CE5-D160876C5847 http://bioguid.osu.edu/xbiod_concepts/8445

98 Caloteleia rubriclava (Ashmead): Ashmead, 1893: 212, 214 (generic transfer, description, keyed). Caloteleia grenadensis Ashmead, 1896: 798 (original description); Ashmead, 1900: 327 (distribution), syn. n. http://zoobank.org/E41DB9D7-F6E1-49BA-BB67-4C8A37430374 http://bioguid.osu.edu/xbiod_concepts/8437 Ceratoteleia Marlatti (Ashmead): Kieffer, 1908: 121 (generic transfer, emendation). Ceratoteleia grenadensis (Ashmead): Kieffer, 1908: 121 (generic transfer); Kieffer, 1926: 501, 505 (description, keyed). Ceratoteleia rubriclava (Ashmead): Kieffer, 1908: 121 (generic transfer); Kieffer, 1926: 501, 504 (description, keyed). Prosanteris nigriceps (Ashmead): Kieffer, 1908: 136 (generic transfer); Kieffer, 1926: 437, 438 (description, keyed). Caloteleia rubriclavus (Ashmead): Brues, 1908: 33 (emendation). Prosanteris (Prosanteris) nigriceps (Ashmead): Kieffer, 1910b: 87 (subgeneric assignment). Caloteleia marlattii (Ashmead): Brues, 1916: 554 (description); Masner & Muesebeck, 1968: 33 (lectotype designation). Macroteleia ruskini Girault, 1920: 179 (original description); Muesebeck, 1958: 93 [junior synonym of Ceratoteleia marlatti (Ashmead)], syn. n. http://zoobank.org/94B4FD7F-E840-437B-8E2B-46F87BE377DF http://bioguid.osu.edu/xbiod_concepts/8447 Ceratoteleia marlattii (Ashmead): Kieffer, 1926: 501, 504 (description, keyed). Calotelea grenadensis (Ashmead): Masner, 1965: 70 (type information). Calotelea nigriceps (Ashmead): Masner & Muesebeck, 1968: 33 (type information). Calotelea rubriclava (Ashmead): Masner & Muesebeck, 1968: 33 (type information). Calotelea ruskini (Girault): Masner & Muesebeck, 1968: 33 (type information, generic transfer).

Description. Body length of female: 1.80–2.55 mm (n=20). Body length of male: 1.60–2.35mm (n=20). Color of head: black throughout; brown throughout. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, short setae. IOS/EH: IOS slightly less than EH; IOS slightly greater than EH.

99 Sculpture of ventrolateral frons: granulate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter; greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Form of male antennal flagellomeres: filiform, A11 approximately 2.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: orange throughout; black throughout; variably orange to pale brown. Color of mesosoma in male: orange throughout; variably orange to pale brown; black throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: rugose throughout. Sculpture of netrion: rugose. Notaulus: percurrent. Sculpture of mesoscutum: granulate; densely punctate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior somewhat rounded or straight, approximately 3.0x wider than long. Sculpture of metascutellum in female: rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: densely punctate. Sculpture of ventral metapleural area: smooth dorsally, densely punctate ventrally. Color of legs: orange throughout; pale brown. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: short, decumbent, hardly exceeding the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as r-rs. Color of metasoma in female: black; orange throughout; variably orange to pale brown. Color of metasoma in male: orange throughout; variably orange to pale brown; black throughout. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: granulate; rugose. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially; reaching posterior margin of T2. Sculpture of T3: smooth; largely

100 smooth with submedian longitudinal striae; longitudinally striate throughout. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: smooth to coriaceous. Diagnosis. This species is easily identified by the combination of the following characters: eye bare; occipital carina complete medially; mesopleural carina absent; mesepisternum below mesopleural depression densely punctate; ventral metapleural area densely punctate ventrally; R1 approximately as long as r-rs. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=4169] Material Examined. Holotype, female, C. grenadensis: GRENADA: Saint George Parish, leeward side, Mount Gay Estate, no date, H. H. Smith, B.M. TYPE HYM. 9.470 (deposited in BMNH). Lectotype, male, C. Marlattii: UNITED STATES: Riley Co., V, Marlatt, USNMENT00989024 (deposited in USNM). Syntype, male, A. nigriceps: UNITED STATES: VA, Arlington Co., Arlington, no date, USNMENT00989028 (deposited in USNM). Holotype, female, A. rubriclavus: UNITED STATES: FL, Duval Co., Jacksonville, ocean beach, Jacksonville Beach (San Pablo), no date, Ashmead, USNMENT00989042 (deposited in USNM). Holotype, female, M. ruskini: UNITED STATES: IL, no date, USNMENT00989079 (deposited in USNM). Other material: (1106 females, 610 males) ARGENTINA: 11 females, 2 males, OSUC 534125–534131, 534437 (CNCI); OSUC 322989, 63153, 63155 (OSUC); OSUC 577255–577256 (UCRC). BAHAMAS: 2 females, 2 males, OSUC 458330–458331, 458383, 534560 (CNCI). BELIZE: 9 females, 3 males, OSUC 534193–534197, 534203, 534214– 534218, 534222 (CNCI). BOLIVIA: 1 female, OSUC 534164 (CNCI). BONAIRE: 2 females, OSUC 458377, 458382 (CNCI). BRAZIL: 48 females, 15 males, OSUC 534527, 553607, 557180–557181, 557197, 557220, 557246, 557248, 557282–557283, 557337, 557342, 557349, 557361, 557375 (CNCI); OSUC 111348, 111698–111700, 111704–111705, 111714, 111810, 111938, 111957, 112073, 112184–112185, 112189–112190, 112194, 112792, 112797, 113088, 11925, 130662, 130709, 130719, 130755, 130849, 131119, 131431, 132407, 132409, 132411, 132413, 132420–132421, 132433, 132446, 133028, 133046, 133050, 133067, 133130, 134362, 137997, 138091, 232034, 367498, 374725, 413, 583202 (OSUC). CANADA: 156 females, 68 males, OSUC 531695–531708, 531710–531720, 531725, 531728–531733, 531802–531865, 531867–531872, 531874–531917, 532076, 532103, 532123–532129, 534491–534493, 534495– 534498, 534500, 534504–534507, 534509–534512, 554065–554117 (CNCI). CHILE: 16 females, OSUC 458265, 534444–534445, 534447, 534449, 534451–534452, 534454, 534456–

101 534458 (CNCI); OSUC 441184–441187 (INHS); OSUC 576993 (OSUC). COLOMBIA: 2 females, 3 males, OSUC 557535, 557537–557539, 557550 (CNCI). COSTA RICA: 24 females, 3 males, OSUC 232068–232069, 532463, 532468, 532494, 532537, 532539, 532569, 532656– 532658, 532664, 532711–532716, 532726, 532765–532766, 532769, 532777–532778, 532792, 532925 (CNCI); OSUC 245767 (OSUC). CUBA: 20 females, 3 males, OSUC 458199, 458201– 458202, 458204–458205, 458207–458211, 458367–458375, 458378, 534558, 534597–534598 (CNCI). DOMINICAN REPUBLIC: 38 females, CMNH-486,514, 490,550, 490,915 (CMNH); OSUC 458178, 458180–458182, 458184, 458186, 458188–458190, 458192, 458194, 458329, 458358–458366, 458376, 458381, 534362–534371, 534374, 534381 (CNCI). ECUADOR: 15 females, OSUC 458519, 458521–458523, 458528, 553368, 553431, 553433, 553601–553602, 553624, 553726–553729 (CNCI). FRENCH GUIANA: 4 females, OSUC 546113–546116 (CNCI). GUATEMALA: 2 females, 2 males, OSUC 534430–534433 (CNCI). HAITI: 2 females, OSUC 534548–534549 (CNCI). HONDURAS: 37 females, 2 males, OSUC 534145 (CNCI); OSUC 399364–399366, 399368–399372, 399374–399377, 399379–399382, 399386– 399387, 399391, 399393, 399395, 399397–399399, 399401–399404, 399407, 399409, 401745, 410436–410438, 410441, 410444–410445, 410448 (MZLU). JAMAICA: 6 females, OSUC 458196–458197, 458379–458380, 534617–534618 (CNCI). KENYA: 1 female, OSUC 56760 (OSUC). MEXICO: 62 females, 44 males, OSUC 531739, 533998–534000, 534004–534012, 534014, 534024, 534464, 534472–534475, 534490 (CNCI); OSUC 410663–410680, 411743, 414931–414932, 49274, 49276, 55926, 583141–583201 (OSUC). NICARAGUA: 2 females, OSUC 534556–534557 (CNCI). PANAMA: 16 females, OSUC 553860–553871, 553873– 553874, 553913–553914 (CNCI). PARAGUAY: 53 females, 8 males, OSUC 150605, 150607, 228804–228807, 276658–276659, 278828, 322986, 322994, 322996–322997, 323036–323037, 323039–323044, 323046–323047, 323049–323053, 323056–323057, 323061–323066, 323068– 323070, 323072–323074, 363636, 363708–363710, 363713, 373987–373997, 534682, 534685, 577154 (OSUC). PERU: 2 females, 1 male, OSUC 554022 (CNCI); OSUC 583203–583204 (OSUC). SAINT LUCIA: 1 female, OSUC 534550 (CNCI). SOUTH AFRICA: 2 males, OSUC 223550–223551 (OSUC). SURINAME: 1 female, OSUC 534569 (CNCI). TRINIDAD AND TOBAGO: 66 females, 1 male, OSUC 545994–545996, 545998, 546000, 546004– 546011, 546014–546022, 546034–546049, 546053–546056, 546059, 546061–546076, 546078– 546081, 546083, 546087, 546091–546092 (CNCI). UNITED STATES: 484 females, 451 males,

102 OSUC 531679–531681, 531683–531685, 531687–531694, 531709, 531721–531724, 531738, 531740–531749, 531751–531775, 531781–531785, 531787–531791, 531793–531801, 531866, 531919–531939, 531948–531980, 531982–532003, 532005–532034, 532037–532044, 532046, 532048–532060, 532062–532063, 532065, 532067–532075, 532078–532099, 532101–532102, 532104–532118, 532120, 532122, 532130, 532247–532303, 532305–532311, 532313–532339, 532391–532399, 532402–532457, 534501–534503, 534508, 554118–554150 (CNCI); OSUC 332917 (MEMU); OSUC 130442, 130451, 130453, 130460, 130462, 130465, 130474, 130477, 130479, 130488, 130491, 130497, 130499–130500, 130513–130515, 130544, 130574, 142758– 142787, 142789–142804, 142806, 142808–142809, 142811–142812, 182700, 182725, 207789, 207795–207796, 236917–236918, 236920, 24231, 256432, 256450–256451, 256513–256517, 256536–256538, 256557–256558, 256595, 256634–256644, 256745–256768, 332914–332916, 336798–336804, 336806–336812, 336814–336818, 336966–336986, 336988–337018, 397539– 397562, 397641–397650, 410662, 55872–55879, 55881–55889, 55891–55906, 576987–576989, 577075–577089, 582301–582320, 583121, 583123–583126, 583128–583140, 583315, 62475, 62495, 62498, 62593, 628906, 62893–62894, 62906, 79811–79812 (OSUC); UCFC 0 118 312, 539, 545, 653, 906, 972, UCFC 0 119 004, UCFC 0 119 183, UCFC 0 119 232, UCFC 0 119 419, UCFC 0 119 434, UCFC 0 120 030, UCFC 0 132 372, UCFC 0 133 615, UCFC 0 133 662, UCFC 0 133 711, UCFC 0 133 782, UCFC 0 133 892, UCFC 0 133 948, UCFC 0 134 479, UCFC 0 134 588, UCFC 0 134 768, UCFC 0 134 851, UCFC 0 134 882, UCFC 0 134 902, UCFC 0 134 963, UCFC 0 135 895, UCFC 0 136 236, UCFC 0 136 271, UCFC 0 136 716, UCFC 0 138 064, UCFC 0 138 130, UCFC 0 138 265, UCFC 0 138 491, UCFC 0 138 502, UCFC 0 138 634, UCFC 0 138 639, UCFC 0 138 772, UCFC 0 138 795, UCFC 0 138 873, UCFC 0 138 898, UCFC 0 138 975, UCFC 0 139 191, UCFC 0 139 197, UCFC 0 139 395, UCFC 0 139 500, UCFC 0 139 633, UCFC 0 139 715, UCFC 0 140 509, UCFC 0 141 638, UCFC 0 142 229, UCFC 0 142 314, UCFC 0 142 556, UCFC 0 142 751, UCFC 0 142 811, UCFC 0 142 817, UCFC 0 143 684, UCFC 0 143 743, UCFC 0 144 035, UCFC 0 144 069, UCFC 0 144 169, UCFC 0 144 180, UCFC 0 144 188, UCFC 0 144 293, UCFC 0 144 568, UCFC 0 144 723, UCFC 0 144 755, UCFC 0 144 878, UCFC 0 144 974, UCFC 0 165 080, UCFC 0 165 131, UCFC 0 165 138, UCFC 0 165 192, UCFC 0 165 195, UCFC 0 165 206, UCFC 0 165 253, UCFC 0 165 546, UCFC 0 165 794, UCFC 0 165 803, UCFC 0 165 808, UCFC 0 165 925, UCFC 0 165 945, UCFC 0 165 952, UCFC 0 166 051, UCFC 0 166 425,

103 UCFC 0 166 683, UCFC 0 166 795, UCFC 0 167 268, UCFC 0 284 802, UCFC 0 319 189, UCFC 0 373 784, UCFC 0 373 785, UCFC 0 374 236, UCFC 0 374 613, UCFC 0 380 130, UCFC 0 380 135, UCFC 0 380 143, UCFC 0 380 144, UCFC 0 380 702, UCFC 0 381 255, UCFC 0 381 256, UCFC 0 389 068, UCFC 0 389 069, UCFC 0 389 076, UCFC 0 389 077, UCFC 0 389 101, UCFC 0 389 105, UCFC 0 389 106, UCFC 0 389 109, UCFC 0 390 859, UCFC 0 391 400, UCFC 0 391 900, UCFC 0 391 901, UCFC 0 392 443, UCFC 0 392 444, UCFC 0 392 447, UCFC 0 392 449, UCFC 0 392 834, UCFC 0 393 389, UCFC 0 393 402, UCFC 0 393 403, UCFC 0 393 871, UCFC 0 393 872, UCFC 0 393 873, UCFC 0 393 874, UCFC 0 393 875, UCFC 0 393 876, UCFC 0 393 877, UCFC 0 393 878, UCFC 0 393 879, UCFC 0 394 467, UCFC 0 394 622, UCFC 0 394 623, UCFC 0 394 624, UCFC 0 394 625, UCFC 0 394 847, UCFC 0 394 848, UCFC 0 395 201, UCFC 0 395 204, UCFC 0 395 485, UCFC 0 395 896, UCFC 0 395 985 (UCFC); OSUC 157686–157687, 157689–157691 (UCMC). VENEZUELA: 23 females, OSUC 545831, 545852, 545862–545863, 545869, 545936, 545948–545950, 546124–546126, 557669–557670, 557674–557675, 557678–557679, 557708, 557711 (CNCI); OSUC 48276, 49158, 64554 (OSUC). Comments. This species seems to be a very common species in the New World, and we also have seen additional specimens from South Africa. Masner et al. (2009) suggested that this kind of widespread distribution may result from commercial traffic. Callisceliorubriclavus exhibits variation in color, ranging from orange to black, in the shape of metascutellum, and in the sculpture of T1 horn and T3. Specimens collected from Central America tend to be darker, while the variation in the sculpture of T1 horn and T3 show no clear correlations among the distrituion ranges

Calliscelio rugaChen & Johnson, new species http://zoobank.org/B725C08C-0768-4304-92AC-9004D072357D http://bioguid.osu.edu/xbiod_concepts/385082 Figures 1.38A–1.38F.

Description. Body length of female: 1.42–1.74 mm (n=20). Color of head: orange throughout. Color of antennal clava (A7–A12): A7, A8 brown, A9–A12 white to pale yellow. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: glabrous. IOS/EH: IOS

104 distinctly less than EH. Sculpture of ventrolateral frons: smooth to granulate. Sculpture of frons below median ocellus: smooth to granulate. Sculpture of posterior vertex: rugose. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: largely smooth with small granulate area. Ocular setae: sparse, short. A4 in female: as long as A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: distinctly longer than wide. Color of mesosoma in female: orange throughout; variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: rugose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin rounded, approximately 3.0x wider than long. Sculpture of metascutellum in female: smooth with a longitudinal, median carina. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose; smooth to rugulose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange yellow. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, weakly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: variably orange to pale brown. Horn on T1 in female: absent. Sculpture of posterior margin of T1 in female: longitudinally striate throughout; striate rugose. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially; present on the anterior margin of T2. Sculpture of T3: smooth. Shape of T6 in female: short, wider than long. Sculpture of S3: largely smooth with sparse and fine punctures. Diagnosis. This species is most similar to C. extenuatus and C. suni in size and habitus. It can be separated from C. extenuatus by the interrupted occipital carina and the elongate A6, and from C. suni by the hairy compound eyes. Etymology. The epithet is used as a noun in apposition derived from the Latin word for wrinkle, in reference to the rugose posterior vertex.

105 Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=385082] Material Examined. Holotype, female: PANAMA: Chiriquí Prov., 1220m, 08°39'N 82°12'W, La Suiza Farm, 16.VI.2000, flight intercept trap, H. Howden, OSUC 553881 (deposited in CNCI). Paratypes: (31 females) ECUADOR: 1 female, OSUC 553619 (CNCI). PANAMA: 29 females, OSUC 553758–553760, 553764, 553793, 553800–553803, 553825, 553827, 553829–553830, 553832, 553845, 553847, 553878–553880, 553882–553889, 553936– 553937 (CNCI). PUERTO RICO: 1 female, OSUC 534561 (CNCI).

Calliscelio rugicoxaChen & Masner, new species http://zoobank.org/ED46B60A-A31E-4165-AD10-C93BFA562781 http://bioguid.osu.edu/xbiod_concepts/362053 Figures 1.39A–1.39F.

Description. Body length of female: 3.16 mm (n=1). Color of head: pale brown. Color of antennal clava (A7–A12): black. Shape of head: subglobose. Central keel of frons: present. Setation of upper frons: with dense, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate to finely punctate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, strongly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate. Ocular setae: dense, long. A4 in female: as long as A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: distinctly longer than wide. Color of mesosoma in female: pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth dorsally, rugulose ventrally. Sculpture of netrion: rugose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: rugose. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: densely punctate. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: smooth with a longitudinal, median carina. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose. Median keels on propodeum in female: present.

106 Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: rugose anteriorly, smooth posteriorly. Sculpture of ventral metapleural area: rugose. Color of legs: pale brown. Sculpture of hind coxa: rugose. Color of fore wing: hyaline. Rs+M: nebulose, weakly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: variably yellow to pale brown. Horn on T1 in female: weakly developed. Sculpture of T1 horn dorsally: rugose. Sculpture of posterior margin of T1 in female: striate rugose. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: largely smooth with submedian longitudinal striae. Shape of T6 in female: distinctly elongate, 2.0x longer than wide. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. bidens but can be distinguished by its rugose hind coxa and rugose T1 horn in the female. Etymology. The epithet refers to the rugose coxa in this species and is intended to be used as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=362053] Material Examined. Holotype, female: COLOMBIA: Valle del Cauca Dept., 650m, 03°26'N 76°48'W, Farallones de Cali Natural National Park, 8.V–19.VI.2001, Malaise trap, S. Sarria, OSUC 458325 (deposited in CNCI). Comments. We generally avoided describing two or more new species based on single specimens when they were collected at the same locality and time, which is the case for C. rugicoxa and C. bidens, but these two species are easily distinguished from each other (see diagnoses of the two species), and we are convinced they are two different species.

Calliscelio sfinaChen & Johnson, new species http://zoobank.org/17D52C03-F10C-4F1C-A73C-0C1C7FE5684C http://bioguid.osu.edu/xbiod_concepts/363178 Figures 1.40A–1.40F.

Description. Body length of female: 1.78–2.97 mm (n=20). Body length of male: 1.67–2.53mm (n=20). Color of head: orange throughout. Color of antennal clava (A7–A12): dark brown to

107 black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: transversely rugulose to granulate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate. Ocular setae: absent. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Form of male antennal flagellomeres: filiform, A11 approximately 2.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: orange throughout. Color of mesosoma in male: orange throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin rounded, approximately 2.5x wider than long. Sculpture of metascutellum in female: rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: rugose. Color of legs: orange throughout; pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, weakly pigmented; spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: orange throughout; yellow with variable pale brown patches. Color of metasoma in male: orange throughout; variably orange to pale brown. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: with V-shaped striae. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth medially, longitudinally striate laterally; longitudinally

108 striate throughout. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: densely punctate medially, longitudinally striate laterally. Diagnosis. This species is most similar to C. carinigena and C. crater in color, size and habitus but it can be distinguished by the absence of the hyperoccipital carina (hyperoccipital carina are absent in C. carinigena and C. crater). Etymology. The specific epithet is Greek for wedge and should be treated as a noun in apposition. It refers to the "wedge-like" horn on T1 in the female. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=363178] Material Examined. Holotype, female: BRAZIL: ES, Duas Bocas Biological Reserve, pt.8, 20°16'21"S 40°28'40"W, Cariacica, 30.IV–1.V.2005, yellow pan trap, A. P. Aguiar et al., OSUC 376920 (deposited in MZSP). Paratypes: (194 females, 236 males) BOLIVIA: 2 females, OSUC 534160, 534182 (CNCI). BRAZIL: 172 females, 226 males, OSUC 534518, 534525- 534526, 557200, 557205-557207 (CNCI); OSUC 577145, 577296, 577299, 577301, 577303, 577369-577370, 577378 (MNHN); OSUC 127850, 150156, 150159, 150785, 150795-150796, 150799, 150803, 150836-150838, 150920, 150957-150958, 150960, 150963, 150980-150983, 150986-150987, 151646, 322521-322524, 322528, 322987, 322999-323000, 323100, 323261- 323263, 323267-323268, 323272, 323280, 323282, 323287, 323292-323295, 323298, 323302, 323305-323307, 323309-323310, 323317, 323323, 323325, 323329-323331, 323337, 323513, 323518-323519, 323521, 323527-323528, 323531-323532, 323538, 323542, 323544, 323546- 323547, 323549, 323551, 323558-323560, 323563, 323566, 323915-323917, 363868, 363881- 363882, 363888, 366644, 366646, 366654-366655, 366658, 372529, 372535, 375247, 375251, 375253, 375263, 375269, 375289, 375291-375292, 375294, 375298, 375302, 376549, 376554, 376917-376919, 376943, 376955, 376962-376963, 381087-381088, 427461, 433802, 433804, 433808, 433810-433811, 433828, 433831, 463311-463312, 576981, 577014, 577032-577033, 577037-577038, 577060-577061, 577070, 577090, 577094, 577096, 577099, 577101, 577103- 577108, 577116-577121, 577199-577203, 577220, 577224-577227 (MZSP); OSUC 110064, 111443, 111648, 11923, 131098, 131107, 131280, 131320, 131673, 132179, 136374, 136540, 136832, 136847, 136857, 136867, 136882, 136934, 136937, 137137, 137182, 137237, 137377, 137430, 137433, 137583, 137599, 137698, 137708, 137885, 138539, 138781, 138784, 147789, 147791-147792, 151400, 151551, 151563, 235891, 235955, 235957-235959, 236824, 236826- 236827, 237004, 322553, 323515, 323517, 347615, 357224-357225, 357246, 357255-357256,

109 357271, 357314, 357321-357322, 357324, 358792, 358940, 358975-358978, 358985-358986, 362661-362664, 362678, 366001, 367437-367438, 367442-367444, 375299, 376923, 378584, 378586-378590, 40192, 427453, 427456-427458, 433805, 433809, 433812-433817, 433825, 433829-433830, 433833-433836, 433838-433841, 433843, 433846, 433848, 435143, 436603, 477161-477162, 510908, 55933, 55938-55940, 55942, 55944, 55954, 576994-576999, 577211, 577272-577275, 577287, 583210-583212, 583226-583227, 583230-583232, 583236, 583290- 583292, 583296, 61314-61315, 61317, 61322, 61335, 61341, 61357-61358, 61364, 61366, 61370, 61375-61376, 61380, 61383, 61407-61408, 61411, 61425, 61430, 61436, 61441, 61443, 61451-61452, 61460-61461, 61463, 61473, 61476, 61502, 61505, 61511, 61521-61522, 61527- 61528, 61535, 61540, 61553, 61558, 63282, 63284, 63291, 63301, 63306, 63322, 63325, 63333, 63348, 63352, 63354, 63381, 63387, 63391-63392, 63401, 63419, 63422, 63428, 63446, 63451, 63491, 63516, 63521, 63565, 63579, 63615, 63663 (OSUC). COLOMBIA: 11 females, 2 males, OSUC 170496, 178186, 178198, 193393, 76994 (IAVH); OSUC 170505, 178020, 178168, 182297, 188679, 188946, 188949, 269347 (OSUC).ECUADOR: 2 females, 4 males, OSUC 458483 (CNCI); OSUC 534677, 534680–534681 (MAIC); OSUC 534669, 534676 (OSUC). FRENCH GUIANA: 1 female, OSUC 534552 (CNCI). GUYANA: 2 females, OSUC 534261– 534262 (CNCI). PARAGUAY: 1 male, OSUC 278677 (OSUC). PERU: 3 females, 1 male, OSUC 323939, 323944, 323946, 323948 (OSUC). TRINIDAD AND TOBAGO: 1 female, OSUC 546026 (CNCI). UNITED STATES: 1 male, OSUC 61320 (OSUC). VENEZUELA: 1 male, OSUC 367474 (USNM).

Calliscelio storea Chen & Johnson, new species http://zoobank.org/ED068E67-CE26-42A0-8DD1-26FD78DD0C72 http://bioguid.osu.edu/xbiod_concepts/384799 Figures 1.2B, 1.41A–1.41F.

Description. Body length of female: 1.76–2.15 mm (n=20). Color of head: black throughout; variably brown to black. Color of antennal clava (A7–A12): black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent.

110 Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate. Ocular setae: sparse, short. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: as long as wide. Color of mesosoma in female: orange throughout; variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth anteriorly, granulate posteriorly. Sculpture of netrion: rugose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: rugose. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugose. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: largely smooth with a row of foveae along mesopleural carina. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: variably orange to pale brown. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: rugulose. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Development of longitudinal striae on T2 in female: present on the anterior margin of T2. Sculpture of T3: smooth. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. bisulcatus and C. virga in the hairy compound eyes, size and habitus. It can be separated from C. bisulcatus by the granulate mesoscutellum and the broad and short metascutellum, and from C. virga by the rugose metascutellum and the rugulose T1 horn in the female. Etymology. The epithet is used as a noun in apposition derived from the Latin word for mat or rug, in reference to the rugulose T1 horn in female. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384799]

111 Material Examined. Holotype, female: FRENCH GUIANA: Cayenne Arrond., 04°33.562'N 52°12.425'W, Kaw Mountains, 3.III–27.III.2007, Malaise trap, K. Sarv, OSUC 546117 (deposited in CNCI). Paratypes: (30 females) BOLIVIA: 1 female, OSUC 534184 (CNCI). BRAZIL: 4 females, OSUC 534619–534620 (CNCI); OSUC 12187, 374720 (OSUC). ECUADOR: 11 females, OSUC 458517, 458520, 458524–458527, 534244, 553400, 553623, 553648, 553716 (CNCI). FRENCH GUIANA: 3 females, OSUC 546118, 546120, 546123 (CNCI). MEXICO: 1 female, OSUC 534622 (CNCI). SURINAME: 1 female, OSUC 534580 (CNCI). VENEZUELA: 9 females, OSUC 545882–545883, 545885–545886, 545889–545890, 545894, 545896, 545941 (CNCI).

Calliscelio suniChen & Johnson, new species http://zoobank.org/7EAA3B0E-69F9-4155-AEE1-58042D054D4A http://bioguid.osu.edu/xbiod_concepts/364057 Figures 1.42A–1.42F.

Description. Body length of female: 1.80–2.18 mm (n=20). Body length of male: 1.70–2.16mm (n=20). Color of head: brown throughout; yellow throughout; orange throughout. Color of antennal clava (A7–A12): A7–A9 brown, A10–A12 white; A7–A9 brown, A10–A12 yellow; brown. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth. Sculpture of frons below median ocellus: smooth. Sculpture of posterior vertex: smooth; granulate; transversely striate; granulate to rugulose above hyperoccipital carina, smooth below. Hyperoccipital carina: absent; present. Occipital carina medially: interrupted. Length of OOL: greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: as long as A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: as long as wide. Form of male antennal flagellomeres: filiform, A11 approximately 4.0x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: orange throughout; yellow throughout. Color of mesosoma in male: orange throughout; variably orange to pale brown; yellow throughout. Sculpture of

112 dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: smooth with sparse punctures; granulate. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: anterior half granulate, posterior half smooth. Shape of metascutellum: posterior marging rounded, approximately 3.0x wider than long. Sculpture of metascutellum in female: smooth; rugose. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose; smooth to rugulose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange throughout; pale yellow throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, weakly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: variably orange to pale brown; yellow throughout. Color of metasoma in male: variably orange to pale brown; brown throughout. Horn on T1 in female: absent. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching the middle of T2 medially; present on the anterior margin of T2. Sculpture of T3: smooth. Shape of T6 in female: short, wider than long or as long as wide. Sculpture of S3: smooth. Diagnosis. This species is similar to C. absum, C. extenuatus, C. minutia and C. ruga in size and habitus. It can be separated from C. absum by the percurrent notaulus, from C. extenuatus and C. ruga by the bare compound eyes and from C. minutia by the smooth frons and the entirely hyaline fore wing. Etymology. The epithet is derived from the name of a famous character in the Chinese classical novel Journey to the West, Sun Wukong, who knows 72 transformations. It refers the remarkable polymorphism of this species and should be treated as a genitive noun. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=364057]

113 Material Examined. Holotype, female: COLOMBIA: Chocó Dept., visitor's center, M.815, 2m, 06°01'N 77°20'W, Utría Natural National Park, 28.IX–15.X.2000, Malaise trap, J. Perez, OSUC 182752 (deposited in IAVH). Paratypes: (1756 females, 235 males) BOLIVIA: 19 females, OSUC 534033, 534045–534047, 534060–534062, 534147–534148, 534152–534155, 534158, 534162–534163, 534183, 534188–534189 (CNCI). BRAZIL: 1297 females, 41 males, OSUC 534534, 534542, 557198-557199, 557203, 557362, 557376-557377 (CNCI); OSUC 251769-251773, 252075-252081, 252083-252085, 252088, 252090-252091, 252093, 252095- 252096, 252098-252104, 252107-252108, 252110, 252112-252114, 252116-252126, 252128- 252129, 252131-252136, 252138, 252140, 252142-252144, 252146-252147, 322153, 322372- 322376, 322378-322380, 322382, 322384, 322533, 322540, 322544, 322549, 322667-322669, 322672-322675, 323006, 323008, 323079, 323082, 323085, 323552, 323929, 323952-323957, 323960-323961, 326190, 326221, 326223, 326225, 326227, 326229, 326231-326234, 326236- 326243, 326245-326247, 326396, 326398-326403, 326500, 326504, 326506-326520, 326534- 326537, 326539-326542, 326544-326545, 337162-337166, 343651-343653, 346126, 346135, 346141, 346161, 346166-346167, 346176, 346185, 346855, 346872-346873, 346881, 346887, 346948, 346951, 347026, 347031, 347042, 347044, 347061, 347072, 347077, 347091, 347955- 347957, 347959, 347962, 347971-347972, 347977-347978, 347980, 347986-347987, 347989, 347995, 347997-347999, 348002-348003, 348006, 348010, 348012, 348066-348067, 348069, 348076, 348101-348102, 348104, 348106, 348116, 348125, 348134, 348139, 348141, 348144, 348147-348148, 348150, 348153-348154, 348264, 348273, 348275, 348278, 348286, 348336, 348347-348348, 348486, 348503, 348506, 348510-348511, 348633, 348640, 348658, 348665, 352050, 352052-352054, 352057, 352060, 352069, 352080-352082, 352360, 352362-352364, 353037, 353040-353044, 353046, 353421-353425, 353428, 353555-353569, 354762, 354793, 354798-354799, 354806, 354813, 354827-354830, 354871-354872, 354893, 354897, 354904- 354905, 355027-355030, 355035, 355037, 355039-355040, 355170-355177, 361081, 361098- 361102, 361104, 361709-361711, 362559, 362587, 362590, 362596-362605, 363848, 363850- 363855, 366691, 366693-366695, 366697, 366699-366700, 366702-366703, 366706-366710, 366713-366720, 366722-366724, 366781, 366783-366789, 366791, 366809-366812, 366937, 366940-366942, 366946-366952, 366967-366973, 366975-366982, 366986-366989, 368048- 368059, 368061, 368411-368412, 368421, 368574-368579, 370900-370904, 370909-370914, 370916-370919, 370922, 370925-370928, 371786, 371815-371816, 371831-371832, 371834-

114 371835, 371853-371854, 372545-372548, 374556, 374558-374559, 374562, 374568-374574, 374576-374585, 374588-374590, 374594, 374622, 374631-374636, 374715-374716, 374733- 374736, 374738, 374743-374748, 376485, 376487, 376489-376491, 376493-376497, 376501- 376504, 376506-376508, 376510-376517, 376525-376526, 376529, 376532-376537, 376539, 376543-376544, 376546-376548, 376930, 376935, 376941-376942, 376967-376975, 376977, 376980, 376983-376984, 377492-377493, 378017-378018, 378020, 378023-378025, 378027- 378031, 378036-378040, 378042-378044, 378046, 378048-378051, 378053, 378056-378059, 378067, 378071, 378073-378079, 378081-378084, 378086-378088, 378601-378602, 378964- 378966, 378968, 378970-378971, 378974, 378978, 378980-378981, 380190-380192 (MZSP); OSUC 235892, 283519-283520, 322383, 322534, 323962-323963, 323965-323969, 323971, 323975-323982, 343650, 343654-343671, 343675-343676, 344391, 344393, 344406, 344411, 344415, 344477, 344486, 344493-344494, 344499-344501, 344511, 344516, 344518, 344526- 344528, 344533-344535, 344545, 345071, 345073, 345075, 345077, 345081, 345083, 345086- 345088, 345090, 345094, 345109, 345117-345118, 345124, 345213, 345217, 345225-345226, 345232, 345250, 345252, 345254, 345257, 345302, 345304, 345310, 345320, 345324, 345329, 345334, 345347, 345421, 345425, 345434, 345438, 345441, 345458, 345467, 345476, 345487, 345536, 345540-345543, 345549, 345553, 345561, 345593, 345859-345860, 345871, 346452- 346454, 346699, 346719, 346724, 346727, 346736-346737, 346739, 346747, 346780-346781, 346791, 346803, 346807, 346809, 346812-346813, 346820, 346823, 346825-346827, 346994, 346998, 347001-347002, 347008, 347010, 347020, 347203, 347209, 347219, 347225, 347228, 347232, 347235, 347239, 347241-347243, 347255-347256, 347269, 347272, 347288, 347388- 347389, 347391, 347393, 347405, 347410, 347413, 347425, 347428-347429, 347433, 347439, 347448-347449, 347473-347474, 347480, 347484-347485, 347494-347495, 347498, 347510, 347512, 347516, 347528-347529, 347531, 347534, 347536, 347538, 347540, 347550, 347552, 347554, 347648, 347651, 347661, 347675-347677, 347680, 347683, 347685, 347735-347736, 347739, 347741, 347752, 347755-347756, 347773-347776, 347842, 347847, 347854, 347876, 347879-347880, 347882-347884, 347886, 347890, 347895, 347898, 347900, 347904, 347910, 348155, 348160, 348958-348959, 348967, 348973, 349002, 349012, 349016, 349020, 349105, 349112-349118, 349120, 349123, 349127, 349141-349144, 349148, 349202-349204, 349207- 349210, 349358-349365, 349369, 349371-349375, 349479-349482, 349484-349488, 349493, 349497, 349502, 349507-349508, 349514, 349516, 349638, 349642-349645, 349647-349648,

115 349650-349651, 349654, 349660-349661, 349664, 349666, 349669, 349673-349674, 349676, 349681, 349684, 349687, 349690, 349694, 349697, 349710, 349712-349713, 349722, 349755, 349757-349762, 349764-349766, 349768-349770, 349772, 349774-349775, 349777-349778, 351244, 351246, 351252, 351262, 351264, 351266, 351269, 351278-351279, 351287, 351294, 351298, 351307, 351313-351314, 351327-351328, 351330-351331, 351397-351398, 351400- 351401, 351404-351405, 351408, 351423, 351450, 351453, 351459-351460, 351465, 351475, 351478-351479, 351499, 351502, 351504, 351523-351524, 351626-351627, 351629, 351674- 351675, 351703, 351808, 351810-351814, 351816, 351820-351822, 351825, 351827-351828, 351830, 351834-351835, 351848, 351854, 351856, 351858-351861, 351865, 351896-351898, 351900-351902, 351905, 352064, 352070, 352072-352075, 352077, 352079, 352361, 352803- 352805, 352807-352809, 352811, 352813, 353178-353180, 353182-353186, 353626, 353645- 353647, 353658-353660, 353668-353669, 353708, 353722-353724, 353728-353730, 353733- 353738, 353740-353744, 353747-353748, 353752, 353755-353756, 353934-353938, 353940, 353943-353944, 353961-353962, 353967, 354040, 354054, 354059, 354092, 354100, 354102, 354111, 354113, 354716-354717, 354720-354721, 354723-354724, 354747, 354749, 355318, 357018-357020, 357025, 357051, 357057-357059, 359048, 366534, 371787-371814, 371828- 371830, 371838, 371845-371848, 373754-373756, 373758-373760, 373763-373769, 373771- 373772, 373774-373778, 373780-373783, 375303-375307, 375309, 375311-375314, 375317, 376052-376056, 376058-376062, 376522-376523, 377498-377503, 378016, 378019, 378021- 378022, 378026, 378032, 378034, 378045, 378047, 378060-378062, 378064, 378066, 378068, 378070, 378072, 378593-378594, 378596, 378598-378600, 381086, 427455, 495357-495383, 534646, 534648, 534650, 534692-534694, 55907-55908, 55948, 577000-577004, 577017, 577125-577127, 577129-577133, 577135-577136, 577151-577153, 577158, 577160, 577163- 577169, 577171-577172, 61523, 63534 (OSUC). COLOMBIA: 162 females, 143 males, OSUC 557414-557419, 557428-557443, 557445-557457, 557473-557475, 557477, 557480-557487, 557489-557496, 557498-557501, 557503-557509, 557511-557514, 557516-557517, 557519- 557525, 557527-557534, 557552, 557557, 557565, 557604-557607, 557613, 557617-557618, 557642-557644 (CNCI); OSUC 144159-144163, 144166, 152141, 162504, 162507, 162591, 162602, 162620, 166587, 178016-178017, 178090, 178093, 178160, 178178-178180, 182595, 188680, 188683, 188686, 188937, 189185, 189203, 189210, 189221, 189295-189296, 191097, 191109-191111, 191146-191147, 192357, 193292, 193295-193296, 193347-193348, 193401,

116 193425, 193543, 193548-193549, 193564, 193595, 193698-193699, 193845, 193861-193862, 193865, 193875, 193877-193878, 193938, 202076-202077, 202080, 231810, 231824, 231827, 232299, 232301, 249900-249901, 253459, 253462, 259752-259753, 259761, 259765, 262545, 262601, 262606-262608, 262941, 262948, 262953-262954, 267808, 268909-268910, 269214- 269215, 269218-269220, 269222, 272081, 272089, 273458, 275810, 276045, 276185-276186, 276243, 279661, 279841, 279905, 279913, 279918, 279924-279925, 279930, 280113, 280180, 280202, 363598, 372636, 377412-377414, 377417, 76993, 76995 (IAVH); OSUC 143971, 143974, 182487, 182741, 182757-182759, 188550, 188625, 188677, 188940, 188942-188943, 188960-188961, 189213, 189215, 189218-189220, 189266, 189269, 189274, 191107, 191151, 191182, 191200, 191298, 191304, 191310, 191314, 191317, 191364, 191367, 191372, 191375, 191385, 191388-191389, 193142, 193145, 193158-193159, 193177, 193194, 193800, 193813, 193846, 253460, 259756, 267809, 269213, 269221, 269223, 269352-269353, 269357, 269430, 269432, 269436, 269440-269443, 269481, 269937-269940, 274570, 274966, 274972, 275026, 279906-279907, 279910, 279917, 280196, 77001, 77005, 77008-77009, 77011 (OSUC). COSTA RICA: 20 females, 1 male, OSUC 532461, 532476, 532495, 532505, 532556, 532575, 532614, 532695–532696, 532709, 532723, 532728, 532730, 532753, 532773, 532786–532787, 532789, 532796, 532831, 532833 (CNCI). ECUADOR: 98 females, 14 males, OSUC 458486, 458489–458490, 458504, 458507, 458512–458513, 458534, 458545, 534226, 534248, 553233– 553236, 553238–553241, 553244–553245, 553248–553254, 553256–553261, 553351–553353, 553355–553365, 553378–553380, 553388, 553402, 553414, 553418–553426, 553501, 553543, 553550, 553552, 553554–553556, 553560, 553572, 553575–553578, 553586–553589, 553603– 553604, 553608–553616, 553654–553655, 553684, 553711, 553713–553715, 553719–553725, 553733–553735, 553745, 577336 (CNCI); OSUC 401740 (MZLU); OSUC 534662 (OSUC). FRENCH GUIANA: 14 females, 2 males, OSUC 458385–458387, 458390, 458409, 458414– 458416, 458422, 458426, 458428–458429, 458471, 534551, 546109, 546130 (CNCI). GUYANA: 9 females, OSUC 534264–534265, 534267–534273 (CNCI). MEXICO: 1 female, OSUC 557254 (CNCI). PANAMA: 78 females, 17 males, OSUC 534069–534081, 534087, 534090, 534092–534096, 534098, 534103–534106, 553750–553751, 553755, 553776–553781, 553785–553792, 553794–553799, 553807, 553811–553813, 553817–553818, 553821, 553824, 553828, 553831, 553833–553835, 553837–553842, 553848–553858, 553890–553891, 553900– 553903, 553906, 553908–553909, 553916–553918, 553920–553921, 553923, 553929, 553950

117 (CNCI). PERU: 45 females, 8 males, OSUC 534388, 534407–534412, 534427–534428, 553969, 553971, 553973–553977, 553979–553985, 553987, 553995–553999, 554001, 554003, 554011– 554015, 554017–554020, 554027, 554029, 554031, 554052 (CNCI); OSUC 199544 (FSCA); OSUC 570541–570544 (OSUC); OSUC 228072–228073, 231995–231996 (USNM). PUERTO RICO: 4 females, 2 males, OSUC 343115–343119, 343124 (OSUC). TRINIDAD AND TOBAGO: 5 females, OSUC 534600, 534602–534604, 546001 (CNCI). UNITED STATES: 3 females, 6 males, OSUC 374705–374706, 374708–374714 (OSUC). VENEZUELA: 1 female, 1 male, OSUC 532725 (CNCI); OSUC 367473 (USNM). Comments. This species is well supported by many characters, although the sculpture of the posterior vertex and the color of the female antennal clava is extremely variable. The posterior vertex varies from smooth to transversely striate. The hyperoccipital carina also can be present or absent. The female antennal clava varies from entirely brown to having the last two or three segments white. These variations are gradual among specimens. Therefore we consider them as intraspecific rather than interspecific.

Calliscelio telumChen & Johnson, new species http://zoobank.org/390805B1-95C4-48E8-8A4A-AAC99C389894 http://bioguid.osu.edu/xbiod_concepts/364059 Figures 1.43A–1.43F.

Description. Body length of female: 2.00–2.89 mm (n=20). Body length of male: 1.99–2.42mm (n=9). Color of head: orange to pale brown; variably brown to black. Color of antennal clava (A7–A12): dark brown to black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with dense, long setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: granulate. Sculpture of frons below median ocellus: granulate. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: granulate. Ocular setae: dense, short. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long.

118 Color of mesosoma in female: orange throughout. Color of mesosoma in male: orange throughout. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth anteriorly, granulate posteriorly. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: granulate. Shape of mesoscutellum: transverse. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: granulate. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: granulate. Sculpture of metascutellum in male: rugose. Dorsal propodeum in female: deeply excavate medially, with lateral propodeal carinae widely separated, running subparallel to accommodate T1 horn. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose with one or two longitudinal keels lateral median keel. Median keels on propodeum in female: absent. Mesopleural carina: present. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: largely smooth, rugose ventrally. Color of legs: orange yellow. Sculpture of hind coxa: smooth. Form of male antennal flagellomeres: filiform, approximately 3.0x greater than width. Length of A5 tyloid in male: longer than 0.5x length of A5. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: distinctly shorter than r-rs. Length of R1: approximately as long as 2.0x length of r-rs. Color of metasoma in female: orange throughout; variably orange to pale brown. Color of metasoma in male: T5–T7 brown to black, otherwise orange. Horn on T1 in female: large and distinct. Sculpture of T1 horn dorsally: rugulose. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: reaching posterior margin of T2. Sculpture of T3: smooth; smooth with longitudinal submedian striae. Shape of T6 in female: distinctly elongate, at least 2.0x longer than wide. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. granulatus in its hairy compound eyes and habitus but can be distinguished by the smooth netrion and rugulose T1 horn. Etymology. The epithet is used as a noun in apposition derived from the Latin word for sword, in reference to the elongate T6 in female. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=364059]

119 Material Examined. Holotype, female: BRAZIL: RO, Rancho Grande Farm, XI.1991, yellow pan trap, S. Passoa, OSUC 48550 (deposited in OSUC). Paratypes: (52 females, 9 males) ARGENTINA: 7 females, OSUC 534132–534136 (CNCI); OSUC 577260–577261 (UCRC). BOLIVIA: 3 females, OSUC 534031, 534180–534181 (CNCI). BRAZIL: 26 females, OSUC 557243, 557338 (CNCI); OSUC 111811, 11920-11922, 11924, 11926, 130907, 131117, 131476, 48560 (MZSP); OSUC 111725, 112704, 130635, 130718, 130750, 130752, 131118, 131188, 132434, 132437, 132572, 374723-374724, 48583 (OSUC). PARAGUAY: 12 females, 9 males, OSUC 534116 (CNCI); OSUC 276697, 276699, 276702, 322995, 322998, 323038, 323048, 323071 (MNHNPY); OSUC 276692-276695, 276698, 276700-276701, 276704, 363635, 434081, 583312-583313 (OSUC).PERU: 1 female, OSUC 323938 (OSUC). URUGUAY: 3 females, OSUC 534607, 534615–534616 (CNCI).

Calliscelio torqueoChen & Johnson, new species http://zoobank.org/97273934-B929-447B-84BE-4E9F0BB09F55 http://bioguid.osu.edu/xbiod_concepts/384798 Figures 1.44A–1.44F.

Description. Body length of female: 1.27–2.54 mm (n=20). Body length of male: 1.39–2.10mm (n=16). Color of head: black throughout; brown throughout; orange throughout. Color of antennal clava (A7–A12): A7–A10 dark brown, A11 and A12 white to pale yellow. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, short setae. IOS/EH: IOS distinctly less than EH. Sculpture of ventrolateral frons: smooth to rugulose. Sculpture of frons below median ocellus: smooth. Sculpture of posterior vertex: smooth. Hyperoccipital carina: absent. Occipital carina medially: interrupted. Length of OOL: greater than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: smooth. Ocular setae: absent. A4 in female: distinctly longer than A3. A5 in female: shorter than A3, distinctly longer than wide. Shape of female A6: distinctly longer than wide. Form of male antennal flagellomeres: filiform, A11 approximately 4.0x longer than wide. Length of A5 tyloid in male: approximately 0.3x length of A5. Color of mesosoma in female: orange throughout; black throughout; variably orange to pale brown. Color of mesosoma in male: orange throughout; variably orange to pale brown;

120 black throughout. Sculpture of dorsal pronotal area: smooth. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: smooth. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: coriaceous; smooth throughout. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: smaller than those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures. Shape of metascutellum: posterior margin somewhat rounded, approximately 2.5x wider than long. Sculpture of metascutellum in female: smooth. Sculpture of metascutellum in male: smooth. Dorsal propodeum in female: not excavate medially, lateral propodeal carinae meeting anteromedially. Sculpture of dorsal propodeum in female: rugose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: fore coxa pale yellow, otherwise variably yellow to pale brown; coxae pale yellow, otherwise orange. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: nebulose, weakly pigmented. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: greater than 3.0x length of r-rs. Color of metasoma in female: orange throughout; variably orange to pale brown. Color of metasoma in male: variably orange to pale brown. Horn on T1 in female: weakly developed. Sculpture of T1 horn dorsally: smooth. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: present on anterior margin of T2 medially, reaching posterior margin of T2 laterally. Sculpture of T3: smooth with longitudinal submedian striae. Shape of T6 in female: short, slightly longer than wide. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. glaber and C. paraglaber in the smooth head and metascutellum. It can be separated from C. glaber by its pigmented Rs+M and smooth small horn, from C. paraglaber by the absence of an occipital carina and T4 is distinctly longer than A3. Etymology. The epithet is used as a noun in apposition derived from the Latin word for torch, in reference to female antennal club color. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=384798]

121 Material Examined. Holotype, female: ECUADOR: Pichincha Prov., 1540m, 00°00'23"N 78°40'36"W, Nanegalito, 27.X–31.X.1999, flight intercept trap, Z. H. Falin, OSUC 553598 (deposited in CNCI). Paratypes: (218 females, 88 males) COLOMBIA: 36 females, 16 males, OSUC 557420–557421, 557471–557472, 557476, 557478–557479, 557488, 557497, 557502, 557510, 557515, 557518, 557526, 557566, 557608, 557612, 557614–557616 (CNCI); OSUC 188727–188728, 188731, 193127, 193692, 202074–202075, 202079, 202081, 202084, 269216, 279912, 279916, 279919–279923, 279927, 280183, 280203, 280206, 377411, 377415– 377416, 377418–377422, 377425–377426 (OSUC). COSTA RICA: 120 females, 38 males, OSUC 232067, 232070, 532466–532467, 532471–532473, 532475, 532477–532478, 532496, 532498, 532500–532501, 532504, 532511–532515, 532520, 532522, 532531, 532534, 532536, 532538, 532540–532542, 532544–532548, 532550, 532555, 532557, 532561–532562, 532564, 532566, 532571, 532579, 532584, 532595–532596, 532609, 532615, 532629, 532642, 532645, 532652, 532654–532655, 532659, 532661–532662, 532671, 532674, 532683, 532690, 532703– 532705, 532707, 532727, 532729, 532731, 532733, 532736, 532742–532745, 532747–532748, 532754–532755, 532758–532764, 532768, 532770, 532772, 532776, 532779–532780, 532782, 532793–532795, 532797–532801, 532803–532822, 532826–532830, 532832, 532835–532840, 532918, 532920–532923, 532926–532927, 532931–532934, 534139–534141, 534236 (CNCI); OSUC 244741, 245170, 245172, 245174, 245237–245238, 246300, 246354–246355, 358624, 374055 (OSUC). ECUADOR: 14 females, 8 males, OSUC 534252, 553354, 553475, 553489, 553495, 553531–553536, 553539, 553574, 553597, 553599, 553617, 553667, 553672–553674, 553708–553709 (CNCI). PANAMA: 25 females, 13 males, OSUC 534086, 534088–534089, 534091, 534097, 534099, 553748–553749, 553752–553754, 553761–553763, 553765–553766, 553772, 553783, 553804–553806, 553815, 553819, 553823, 553836, 553843–553844, 553846, 553859, 553875–553876, 553928, 553933–553934, 553941–553943, 553951 (CNCI). VENEZUELA: 23 females, 13 males, OSUC 545830, 545871–545872, 545939–545940, 545955, 545957, 545991–545993, 557657, 557701–557707, 557712 (CNCI); OSUC 146810, 334525, 334545, 334547, 46271, 46275, 46569, 48153, 48213, 48232, 48327, 48481, 48756, 48784, 48789, 63915, 79762 (OSUC).

Calliscelio virgaChen & Johnson, new species http://zoobank.org/D2DB314F-6D8F-44B6-BB3C-912BD3DC0029

122 http://bioguid.osu.edu/xbiod_concepts/364058 Figures 1.45A–1.45F.

Description. Body length of female: 1.68–2.10 mm (n=20). Body length of male: 1.64–2.07mm (n=20). Color of head: brown throughout; variably brown to black. Color of antennal clava (A7– A12): black. Shape of head: subglobose. Central keel of frons: absent. Setation of upper frons: with sparse, long setae. IOS/EH: IOS slightly less than EH. Sculpture of ventrolateral frons: smooth to granulate. Sculpture of frons below median ocellus: largely smooth with sparse fine punctures. Sculpture of posterior vertex: granulate. Hyperoccipital carina: absent. Occipital carina medially: complete, weakly crenulate throughout. Length of OOL: less than 0.5x ocellar diameter. Sculpture of postgena behind outer orbit: largely smooth with small granulate area. Ocular setae: sparse, short. A4 in female: distinctly shorter than A3. A5 in female: shorter than A3, as long as wide. Shape of female A6: distinctly wider than long. Form of male antennal flagellomeres: filiform, A11 approximately 2.0x longer than wide. Length of A5 tyloid in male: greater than 0.5x length of A5. Color of mesosoma in female: orange throughout; variably orange to pale brown. Color of mesosoma in male: variably orange to pale brown. Sculpture of dorsal pronotal area: rugose. Sculpture of lateral pronotal area: smooth throughout. Sculpture of netrion: rugose. Notaulus: percurrent or nearly so. Sculpture of mesoscutum: smooth with sparse punctures; coriaceous. Shape of mesoscutellum: semiellipsoidal. Foveolae of scutoscutellar sulcus between notauli: as large as those along margin of axilla. Sculpture of mesoscutellum: smooth with sparse fine punctures; granulate. Shape of metascutellum: posterior margin straight, approximately 4.0x wider than long. Sculpture of metascutellum in female: smooth. Sculpture of metascutellum in male: smooth. Dorsal propodeum in female: shallowly excavate medially, with lateral propodeal carinae widely separated. Sculpture of dorsal propodeum in female: rugulose. Sculpture of dorsal propodeum in male: rugose. Median keels on propodeum in female: absent. Mesopleural carina: absent. Sculpture of mesepisternum below mesopleural depression: smooth. Sculpture of ventral metapleural area: smooth. Color of legs: orange throughout. Sculpture of hind coxa: smooth. Color of fore wing: hyaline. Rs+M: spectral. Setae on R: long, erect, surpassing the margin of the wing. Length of R: approximately as long as r-rs. Length of R1: approximately as long as r-rs.

123 Color of metasoma in female: orange throughout; variably orange to pale brown. Color of metasoma in male: variably orange to pale brown. Horn on T1 in female: present as a small bulge. Sculpture of T1 horn dorsally: smooth. Sculpture of posterior margin of T1 in female: longitudinally striate throughout. Sculpture of T1 in male: longitudinally striate. Development of longitudinal striae on T2 in female: present on the anterior margin of T2. Sculpture of T3: smooth. Shape of T6 in female: short, wider than long. Sculpture of S3: smooth. Diagnosis. This species is most similar to C. bisulcatus and C. storea in the hairy compound eyes, size and habitus but can be easily recognized by its narrow and smooth metascutellum. Etymology. The specific epithet is Latin for rod or wand and should be treated as a noun in apposition. It refers to the narrow smooth metascutellum of this species. Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=364058] Material Examined. Holotype, female: BOLIVIA: Santa Cruz Dept., Andrés Ibáñez Prov., B-21, pools, 375m, 17°40'S 63°27'W, El Hondo, 14.V–17.V.1997, yellow pan trap, L. Masner, OSUC 534034 (deposited in CNCI). Paratypes: (169 females, 98 males) BRAZIL: 15 females, OSUC 534523, 557298 (CNCI); OSUC 48514, 48520, 55945, 55947, 55952, 583247, 583250, 583252–583255, 583257–583258 (OSUC). CANADA: 8 males, OSUC 531726– 531727, 531734–531737, 532035, 532077 (CNCI). COLOMBIA: 1 female, 1 male, OSUC 557585, 557623 (CNCI). COSTA RICA: 1 female, OSUC 532651 (CNCI). DOMINICAN REPUBLIC: 2 males, OSUC 534361, 534375 (CNCI). ECUADOR: 3 females, OSUC 458516, 458518, 553369 (CNCI). GUATEMALA: 1 male, OSUC 534434 (CNCI). MEXICO: 5 males, OSUC 534018–534020, 534477–534478 (CNCI). PARAGUAY: 3 females, 4 males, OSUC 150470, 534684, 577155, 577191–577192, 577195–577196 (OSUC). SURINAME: 4 females, OSUC 534568, 534572, 534587, 553621 (CNCI). TRINIDAD AND TOBAGO: 2 females, OSUC 546029, 546085 (CNCI). UNITED STATES: 138 females, 76 males, OSUC 531682, 531686, 531750, 531776–531780, 531786, 531792, 531918, 531940–531947, 532061, 532064, 532100, 532119, 532121, 532131–532177, 532180–532246, 532340–532390, 534347–534360 (CNCI); OSUC 142805, 142807, 142810, 207785, 236919, 272939–272940, 576980, 62904, 62907, 62927 (OSUC). VENEZUELA: 2 females, 1 male, OSUC 557676 (CNCI); OSUC 55924–55925 (OSUC).

124

Figure 1.1 A, Calliscelio punctatifronssp. n., female, holotype (OSUC 191124), Antenna. B–C, Calliscelio punctatifronssp. n., male, paratype (OSUC 191125). B, Antenna; C, Tyloid. D, Calliscelio latifrons sp. n., female, holotype (OSUC 323077), Antenna. E–F, Calliscelio latifrons sp. n., male, paratype (OSUC 323075). E, Antenna; F, Tyloid.G, Calliscelio alcoa sp. n., female, holotype (OSUC 458212), Antenna. H–I, Calliscelio alcoa sp. n., male, paratype (OSUC 458222). H, Antenna; I, Tyloid.J, Calliscelio longius sp. n., female, paratype (OSUC 193935), Antenna. K–L, Calliscelio longius sp. n., male, paratype (OSUC 193596). K, Antenna; L, Tyloid.

125

Figure 1.2 Fore wing, dorsal view. A, Calliscelio gatineau sp. n., female, holotype (OSUC 534340). B, Calliscelio storea sp. n., female, holotype (OSUC 546117). C, Calliscelio laticinctus Ashmead, female (OSUC 458242).

126

Figure 1.3 A, Calliscelio absconditumsp. n., female, paratype (OSUC 134791), dorsal habitus. B–F,Calliscelio absconditumsp. n., female, holotype (OSUC 323924). B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

127

Figure 1.4 Calliscelio absum sp. n., female, holotype (OSUC 190716). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

128

Figure 1.5 Calliscelio alcoa sp. n., female, holotype (OSUC 458212). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

129

Figure 1.6 Calliscelio amadoi sp. n., female, holotype (OSUC 193589). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

130

Figure 1.7 Calliscelio armila sp. n., female, holotype (OSUC 458342). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

131

Figure 1.8 Calliscelio bidens sp. n., female, holotype (OSUC 458326). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

132

Figure 1.9 Calliscelio bisulcatus (Kieffer), female, holotype (CAS Type No. 9590). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

133

Figure 1.10 Calliscelio brachys sp. n., female, holotype (OSUC 193349). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

134

Figure 1.11 Calliscelio brevinotaulus sp. n., female, holotype (OSUC 190678). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

135

Figure 1.12 Calliscelio brevitas sp. n., female, holotype (OSUC 190705). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

136

Figure 1.13 Calliscelio carinigena sp. n., female, holotype (OSUC 343907). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

137

Figure 1.14 Calliscelio crater sp. n., female, holotype (OSUC 276803). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

138

Figure 1.15 Calliscelio crena sp. n., female, holotype (OSUC 553462). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

139

Figure 1.16 Calliscelio eboris sp. n., female, holotype (OSUC 193404). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

140

Figure 1.17 Calliscelio extenuatus sp. n., female, holotype (OSUC 48454). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

141

Figure 1.18 Calliscelio flavicauda sp. n., female, holotype (OSUC 553509). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

142

Figure 1.19 Calliscelio foveolatus sp. n., female, holotype (OSUC 532869). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

143

Figure 1.20 Calliscelio gatineau sp. n., female, holotype (OSUC 534340). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

144

Figure 1.21 Calliscelio glaber sp. n., female, holotype (OSUC 458136). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

145

Figure 1.22 Calliscelio granulatus sp. n., female, holotype (OSUC 534599). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

146

Figure 1.23 Calliscelio laticinctus Ashmead, female, syntype (USNMENT01059105). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Specimen labels.

147

Figure 1.24 Calliscelio latifrons sp. n., female, holotype (OSUC 323077). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

148

Figure 1.25 Calliscelio levis sp. n., female, holotype (OSUC 553454). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

149

Figure 1.26 Calliscelio longius sp. n., female, holotype (OSUC 374730). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

150

Figure 1.27 Calliscelio magnificus sp. n., female, holotype (OSUC 458272). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

151

Figure 1.28 Calliscelio migma sp. n., female, holotype (OSUC 380181). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

152

Figure 1.29 Calliscelio minutia sp. n., female, holotype (OSUC 433847). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

153

Figure 1.30 Calliscelio paraglaber sp. n., female, holotype (OSUC 532684). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

154

Figure 1.31 Calliscelio pararemigio sp. n., female, holotype (OSUC 458239). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

155

Figure 1.32 Calliscelio prolixus sp. n., female, holotype (OSUC 557587). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

156

Figure 1.33 Calliscelio punctatifrons sp. n., female, holotype (OSUC 191124). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

157

Figure 1.34 Calliscelio remigio sp. n., female, holotype (OSUC 458296). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

158

Figure 1.35 A–B, Acolus rubriclavus Ashmead, female, holotype (USNMENT00989042). A, Mesosoma and metasoma, lateral view; B, Mesosoma and metasoma, dorsal view. C–F, Anteris nigriceps Ashmead, male, syntype (USNMENT00989028). C, Mesosoma and metasoma, lateral view; D, Mesosoma, lateral view; E, Mesosoma and metasoma, dorsal view; F, Mesosoma, dorsal view.

159

Figure 1.36 Caloteleia grenadensis Ashmead, female, holotype (B.M. TYPE HYM. 9.470). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Specimen labels.

160

Figure 1.37 A–B, Caloteleia marlattii Ashmead, male, lectotype (USNMENT00989024). A, Lateral habitus; B, Dorsal habitus; C–F, Macroteleia ruskini Girault, female, holotype (USNMENT00989079). C, Mesosoma and metasoma, lateral view; D, Mesosoma and metasoma, dorsal view; E, Metasoma, dorsal view; F, Metasoma, ventral view.

161

Figure 1.38 Calliscelio ruga sp. n., female, holotype (OSUC 553881). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

162

Figure 1.39 Calliscelio rugicoxa sp. n., female, holotype (OSUC 458325). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

163

Figure 1.40 Calliscelio sfina sp. n., female, holotype (OSUC 376920). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

164

Figure 1.41 Calliscelio storea sp. n., female, holotype (OSUC 546117). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

165

Figure 1.42 Calliscelio suni sp. n., female, holotype (OSUC 182752). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

166

Figure 1.43 Calliscelio telum sp. n., female, holotype (OSUC 276704). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

167

Figure 1.44 Calliscelio torqueo sp. n., female, holotype (OSUC 553598). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

168

Figure 1.45 Calliscelio virga sp. n., female, holotype (OSUC 534034). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F,Metasoma, dorsal view.

169 Table 1.1 List of morphological terms and links to Hymenoptera Anatomy Ontology database for New World Calliscelio.

A1 http://purl.obolibrary.org/obo/HAO_0000908 A2 http://purl.obolibrary.org/obo/HAO_0000706 A3 http://purl.obolibrary.org/obo/HAO_0001148 A7 http://purl.obolibrary.org/obo/HAO_0001885 A12 http://purl.obolibrary.org/obo/HAO_0001884 antenna http://purl.obolibrary.org/obo/HAO_0000101 antennomere http://purl.obolibrary.org/obo/HAO_0000107 area http://purl.obolibrary.org/obo/HAO_0000146 body http://purl.obolibrary.org/obo/HAO_0000182 carina http://purl.obolibrary.org/obo/HAO_0000188 central keel http://purl.obolibrary.org/obo/HAO_0000109 cpa cervical pronotal area http://purl.obolibrary.org/obo/HAO_0000194 clava http://purl.obolibrary.org/obo/HAO_0000203 clypeus http://purl.obolibrary.org/obo/HAO_0000212 compound eye http://purl.obolibrary.org/obo/HAO_0000217 coxa http://purl.obolibrary.org/obo/HAO_0000228 depression http://purl.obolibrary.org/obo/HAO_0000241 dpa dorsal pronotal area http://purl.obolibrary.org/obo/HAO_0000267 egg http://purl.obolibrary.org/obo/HAO_0000286 epomial carina http://purl.obolibrary.org/obo/HAO_0000307 eye http://purl.obolibrary.org/obo/HAO_0000217 femur http://purl.obolibrary.org/obo/HAO_0000327 fore wing http://purl.obolibrary.org/obo/HAO_0000351 frons http://purl.obolibrary.org/obo/HAO_0001523 gena http://purl.obolibrary.org/obo/HAO_0000371 head http://purl.obolibrary.org/obo/HAO_0000397 hind coxa http://purl.obolibrary.org/obo/HAO_0000587 hind tibia http://purl.obolibrary.org/obo/HAO_0000631 hind wing http://purl.obolibrary.org/obo/HAO_0000400 inner orbit http://purl.obolibrary.org/obo/HAO_0000419 interantennal process http://purl.obolibrary.org/obo/HAO_0000422 lateral lobe of mesoscutum http://purl.obolibrary.org/obo/HAO_0000466 lateral ocellus http://purl.obolibrary.org/obo/HAO_0000481 LOL lateral ocellar line http://purl.obolibrary.org/obo/HAO_0000480 lpa lateral pronotal area http://purl.obolibrary.org/obo/HAO_0000483 malar sulcus http://purl.obolibrary.org/obo/HAO_0000504 mandible http://purl.obolibrary.org/obo/HAO_0000506 margin http://purl.obolibrary.org/obo/HAO_0000510 mesepisternum http://purl.obolibrary.org/obo/HAO_0001872 med mesopleural depression http://purl.obolibrary.org/obo/HAO_0000326 mesopleuron http://purl.obolibrary.org/obo/HAO_0000566

170 Table 1.1. Continued mesoscutellum http://purl.obolibrary.org/obo/HAO_0000574 mesoscutum http://purl.obolibrary.org/obo/HAO_0001490 mesosoma http://purl.obolibrary.org/obo/HAO_0000576 metapleuron http://purl.obolibrary.org/obo/HAO_0000621 metascutellum http://purl.obolibrary.org/obo/HAO_0000625 metasoma http://purl.obolibrary.org/obo/HAO_0000626 midlobe of mesoscutum http://purl.obolibrary.org/obo/HAO_0000520 netrion http://purl.obolibrary.org/obo/HAO_0000644 notauli (notaulus) http://purl.obolibrary.org/obo/HAO_0000647 occipital carina http://purl.obolibrary.org/obo/HAO_0000653 ocellus http://purl.obolibrary.org/obo/HAO_0000661 ot ocellar triangle http://purl.obolibrary.org/obo/HAO_0000430 OOL ocular ocellar line http://purl.obolibrary.org/obo/HAO_0000662 orbit http://purl.obolibrary.org/obo/HAO_0000672 POL posterior ocellar line http://purl.obolibrary.org/obo/HAO_0000759 process http://purl.obolibrary.org/obo/HAO_0000822 propodeum http://purl.obolibrary.org/obo/HAO_0001248 S1 http://purl.obolibrary.org/obo/HAO_0001997 S2 http://purl.obolibrary.org/obo/HAO_0001829 S3 http://purl.obolibrary.org/obo/HAO_0001831 S7 http://purl.obolibrary.org/obo/HAO_0002185 sculpture http://purl.obolibrary.org/obo/HAO_0000913 sternite http://purl.obolibrary.org/obo/HAO_0001654 sulcus http://purl.obolibrary.org/obo/HAO_0000978 T1 http://purl.obolibrary.org/obo/HAO_0000053 T2 http://purl.obolibrary.org/obo/HAO_0000056 T3 http://purl.obolibrary.org/obo/HAO_0000057 T4 http://purl.obolibrary.org/obo/HAO_0000058 T5 http://purl.obolibrary.org/obo/HAO_0000059 T6 http://purl.obolibrary.org/obo/HAO_0000060 T7 http://purl.obolibrary.org/obo/HAO_0000061 tergite http://purl.obolibrary.org/obo/HAO_0001783 tibia http://purl.obolibrary.org/obo/HAO_0001017 tyloid http://purl.obolibrary.org/obo/HAO_0001199 vein http://purl.obolibrary.org/obo/HAO_0001095 vertex http://purl.obolibrary.org/obo/HAO_0001077 vertical epomial carina http://purl.obolibrary.org/obo/HAO_0000307 wing http://purl.obolibrary.org/obo/HAO_0001089

171 Acknowledgments

Thanks to E. Talamas (USNM) for discussion and to L. Musetti, J. Cora, and S. Hemly (OSUC) for critical assistance with specimen handling, software, and databasing. This material is based upon work supported in part by the U.S. National Science Foundation under grant No. DEB-0614764 to N.F. Johnson and A.D. Austin.

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175 Chapter 2: Revision of the World species of the Genus Habroteleia Kieffer (Hymenoptera, Platygastridae, Scelioninae)

Abstract The genus Habroteleia Kieffer is revised. Seven species are recognized, three are redescribed: H. flavipes Kieffer, H. persimilis (Kozlov & Kononova), H. ruficoxa (Kieffer); and four species are described as new: H. mutabilis Chen & Talamas, n. sp., H. salebra Chen & Talamas, n. sp., H. soa Chen & Talamas, n. sp., and H. spinosa Chen & Johnson, n. sp. Four species are treated as junior synonyms of Habroteleia flavipes Kieffer: Chrestoteleia bakeri Kieffer, n. syn., Habroteleia bharatensis Saraswat, n. syn., Habroteleia browni Crawford, n. syn., and Habroteleia kotturensis (Sharma), n. syn. Habroteleia dagavia (Kozlov & Lê), n. syn. is treated as junior synonym of Habroteleia persimilis (Kozlov & Kononova). Baryconus vindhiensis (Sharma), n. comb. is transferred out of Habroteleia Kieffer. Habroteleia impressa (Kieffer) and H. scapularis (Kieffer) remain valid species but their identity and status are unclear.

Introduction The genus Habroteleia was originally described by Kieffer (1905) based on the type species, Habroteleia flavipes Kieffer, collected on the island of Sumatra, Indonesia. Kieffer (1913) later proposed Chrestoteleia for a single species, Chrestoteleia bakeri Kieffer, collected from the Philippines, which was treated by Baltazar (1961) as a junior synonym of Habroteleia. Nine species have since been described from India, Japan and the Philippines. We here provide the first comprehensive treatment of the genus, including examination of type specimens of all species except H. impressa (Kieffer) and H. scapularis (Kieffer), for which we were unable to locate type material. The previously described species of Habroteleia were recorded from the Oriental region, extending from India to Japan, and we here provide records that expand the distribution of Habroteleia to include Madagascar, Papua New Guinea, and the Fijian archipelago. The host of Habroteleia is unknown, but we suspect that it parasitizes orthopteran eggs (large and elongate) based on its elongate body and that Orthoptera suspected as the plesiomorphic host group for the scelionids (Austin et al. 2005).

176 The contributions of the authors are as follows. H.-Y. Chen, E. J. Talamas and N.F. Johnson: character definition, generic concept development, species concept development, imaging, key development, manuscript preparation; L. Masner: character definition, generic concept development, species concept development. The authors of the new species are indicated in the heading of each description.

Materials and methods

This work is based upon specimens in the following collections, with abbreviations used in the text: BPBM, Bernice P. Bishop Museum, Honolulu, HI; CNCI, Canadian National Collection of Insects, Ottawa, Canada; CSCA, California State Collection of Arthropods, Sacramento, CA; FSCA, Florida State Collection of Arthropods, Gainesville, FL; IEBR, Institute of Ecology and Biological Resources, Hanoi, Vietnam; MCSN, Museo Civico de Storia Naturale "Giacomo Doria", Genoa, Italy; MNHN, Muséum National d'Histoire Naturelle, Paris, France; OSUC, C.A. Triplehorn Insect Collection, Ohio State University, Columbus, OH; SCAU, South China Agricultural University, Guangzhou, China; UCDC, R.M. Bohart Museum of Entomology, University of California, Davis, CA; ZIN, Zoological Museum, Academy of Sciences, St. Petersburg, Russia. 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); EH: eye height, length of compound eye measured parallel to dorsoventral midline of head; IOS: interocular space, minimal distance on frons between compound eyes; OD: ocellar diameter, greatest width of ocellus; OOL: ocular ocellar line, shortest distance from inner orbit and outer margin of posterior ocellus (Masner 1980); T1, T2, ... T7: metasomal tergite 1, 2, ... 7; S1, S2, … S7: metasomal sternite 1, 2, … 7. Morphological terminology otherwise generally follows Masner (1980) and Mikó et al. (2007). Morphological terms used in this work were as in the Hymenoptera Anatomy Ontology (Yoder et al. 2010) (Table 2.1). Identifiers (URIs) in the format HAO_XXXXXXX represent concepts in the HAO and are provided to enable readers to confirm their understanding of the concepts being referenced. To learn more about a given concept, including additional images,

177 notes, references and other metadata, use the identifier as a search term at http://glossary.hymao.org or use the identifier as a web-link. In the Material Examined section the metadata for the specimens studied are recorded in an abbreviated format, using unique identifiers (numbers prefixed with “OSUC”, “CASENT”, “FBA”, “MNHN_EY”) 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 can be accessed at the following link, hol.osu.edu, and entering the identifier in the form (note the space between the acronym and the number). 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, www.zoobank.org), and other taxonomic names, where appropriate, have been retrospectively registered. The external hyperlinks are explicitly cited in the endnotes so that users of the printed version of this article have access to the same resources. Data associated with the genus Habroteleia can be accessed at http://hol.osu.edu/index.html?id=488. The generic and species descriptions were generated by a database application, vSysLab (vsyslab.osu.edu), designed to facilitate the production of a taxon by character data matrices, and to integrate those data with the existing taxonomic, media, 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 are indicated by semicolon-separated character states. Images and measurements were produced with multiple systems. Photographs of IEBR specimens were captured with a Canon Rebel 600 camera connected to a Wild M10 microscope with a Fotoprojektiv 2.5x/SLR 10446175 adapter and stacked with the program Zerene Stacker. A scale bar was calibrated for images taken at the maximum magnification of the microscope. The remaining images were produced with Combine ZP and AutoMontage extended-focus software, using a JVC KY-F75U digital camera, Leica Z16 APOA microscope, and 1X objective lens. Images were post-processed with Adobe Photoshop CS3 Extended. A standard set of images is provided for each species: dorsal habitus, lateral habitus, dorsal and lateral views of the

178 head and mesosoma, and anterior view of head. The individual images are archived in Specimage (specimage.osu.edu), the image database at The Ohio State University. Images of primary types of H. ruficoxa and H. persimilis were provided by Agnièle Touret- Alby (MNHN) and Konstantin Samartsev (ZIN), respectively. Images of the primary type of Baryconus vindhiensis, Habroteleia bharatensis and Habroteleia kotturensis were made available by Talamas et al. (2017) and images of Triteleia dagavia were made available by Talamas & Pham (2017), all are used in this publication with permission.

Taxonomy

Habroteleia Kieffer

Habroteleia Kieffer, 1905: 14 (original description. Type: Habroteleia flavipes Kieffer, by monotypy); Kieffer, 1908: 114 (keyed); Brues, 1908: 27, 38 (diagnosis, list of species, keyed); Kieffer, 1910: 63, 69 (description, list of species, keyed); Kieffer, 1913: 220 (description); Kieffer, 1926: 267, 363 (description, keyed, key to species); Muesebeck & Walkley, 1956: 357 (citation of type species); Baltazar, 1961: 395 (synonymy); Baltazar, 1966: 177 (cataloged, catalog of species of the Philippines); Masner, 1976: 10, 26 (description, keyed); Mani & Sharma, 1982: 155, 167 (description, keyed); Johnson, 1992: 398 (cataloged, catalog of world species); Austin & Field, 1997: 24, 68 (structure of ovipositor system, discussion of phylogenetic relationships, genus misplaced in Calliscelionini); Lê, 2000: 31 (keyed); Kononova & Kozlov, 2008: 23, 255 (description, keyed); Chen, Johnson, Masner & Xu, 2013: 11 (keyed). http://zoobank.org/CBFA7C74-68DD-44F2-BE05-AEBD88E6FA8D http://bioguid.osu.edu/xbiod_concepts/488 Chrestoteleia Kieffer, 1913: 388 (original description. Type: Chrestoteleia bakeri Kieffer, by monotypy and original designation. Synonymized by Baltazar (1961)); Kieffer, 1926: 271, 442 (description, keyed, key to species); Muesebeck & Walkley, 1956: 342 (citation of type species); Baltazar, 1961: 395 (junior synonym of Habroteleia Kieffer); Baltazar, 1966: 182 (cataloged, catalog of species of the Philippines). http://zoobank.org/4EA90A05-D50A-42BF-B1C0-852F4B56FCBA

179 http://bioguid.osu.edu/xbiod_concepts/8933 Crestoteleia Kieffer: Kieffer, 1916: 180 (key to new species described from the Philippines, spelling error).

Description. Length 2.18–5.18 mm; body moderately to markedly elongate, robust. Head. Head shape in dorsal view: transverse. Hyperoccipital carina: absent. Occipital carina: present, complete or broadly interrupted medially. Anterior margin of occipital carina: crenulate. OOL: lateral ocellus nearly contiguous with inner orbits, OOL < 0.5 OD; lateral ocellus contiguous with inner orbit. Upper frons: convex, without frontal shelf or carina. Antennal scrobe: broadly convex or conave medially with distinct depression. Sculpture of antennal scrobe: smooth to punctate. Submedian carina: absent. Orbital carina: absent. Inner orbits: diverging ventrally. IOS/EH: IOS distinctly less than EH. Interantennal process: short, often excavate medially. Central keel: present or absent. Antennal foramen: oriented laterally on interantennal process. Facial striae: absent. Malar sulcus: present. Setation of compound eye: absent. Gena: broad, convex, distinctly produced behind eye. Clypeus shape: narrow, slightly convex medially, lateral corners not produced. Anterior (or ventral) margin of clypeus: straight. Anteclypeus: absent. Postclypeus: absent. Labrum: not visible in anterior view. Number of mandibular teeth: 2. Arrangement of mandibular teeth: transverse. Number of maxillary palpomeres: 4. Shape of maxillary palpomeres: cylindrical. Number of labial palpomeres: 2. Antenna. 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. Number of antennomeres with multiporous plate sensilla in female: 5.Arrangement of doubled multiporous plate sensilla on female clava: in longitudinal pairs. Number of antennomeres bearing tyloids in male antenna: 0. Shape of male flagellum: filiform. Mesosoma. Transverse pronotal carina: present anterior to epomial carina, present or absent posterior to epomial carina. Posterior apex of pronotum in dorsal view: straight, bifid apically to articulate with tegula. Epomial carina: present. Anterior face of pronotum: oblique, visible dorsally, short. Lateral face of pronotum: weakly concave below position of dorsal epomial carina. Netrion: present. Netrion shape: moderately wide, open ventrally. Anterior

180 portion of mesoscutum: vertical, flexed ventrally to meet pronotum. Mesoscutum shape: pentagonal, excavate at base of wings. Skaphion: absent. Notauli: present, percurrent. Parapsidal lines: absent. Antero-admedian lines: absent. Transscutal articulation: well-developed, narrow. Shape of mesoscutellum: trapezoidal. Lateral mesoscutellar spine: absent. Median mesoscutellar spine: absent. Axillular spine: absent. Surface of mesoscutellum: convex throughout. Median longitudinal furrow on mesoscutellum: absent; present. Metascutellum: clearly differentiated. Form of metascutellum: transverse. Posterior margin of metascutellum: straight with a small projection medially. Setation of metascutellum: absent. Metapostnotum: not defined externally. Lateral propodeal projection: present. Median propodeal projection: present. Mesopleural carina: present. Mesal course of acetabular carina: not separating fore coxae. Mesopleural pit: present. Posterodorsal corner of mesopleuron: rounded anteriorly. Legs. Number of mesotibial spurs: 1. Number of metatibial spurs: 1. Dorsal surface of metacoxa: smooth; punctate. Shape of metacoxa: cylindrical, ecarinate. Trochantellus: indicated by transverse sulcus on femur. Wings. Wing development of female: macropterous. Wing development of male: macropterous. Tubular veins in fore wing: present. Bulla of fore wing R: absent. Length of marginal vein of fore wing: more than twice as long as stigmal vein. Origin of r-rs in fore wing: arising from marginal vein along costal margin. Basal vein (Rs+M) in fore wing: absent. Development of R vein in hind wing: complete. Metasoma. Number of external metasomal tergites in female: 6. Number of external metasoma sternites in female: 6. Number of external metasomal tergites in male: 8. Number of external metasomal sternites in male: 8. Shape of metasoma: lanceolate. Laterotergites: present, narrow. Laterosternites: present. T1 of female: flat; medially convex as a small hump anteriorly. Relative size of metasomal segments: T3 longest, T2 and T4 subequal in length. Metasomal tergites with basal crenulae: T2. Sublateral carinae on tergites: absent. Median longitudinal carina on metasomal terga: absent. Shape of female T6: flattened. Anterior margin of S1: not produced anteriorly, straight. Felt fields: absent. Ovipositor: Ceratobaeus-type (Austin & Field 1997). Diagnosis. Habroteleia can be separated from other scelionines by the combination of the following characters: epomial carina present; malar and facial striae absent; marginal vein many times longer than stigmal vein; postmarginal vein (R1) absent or rudimentary; propodeum with

181 lateral and median projections; T6 in females strongly depressed dorsoventrally to form a flat triangle; male antenna without tyloid (Chen et al. 2013). The wing venation and large size of Habroteleia make it a relatively easy genus to identify. In all species of Habroteleia the marginal vein is many times longer than the stigmal vein and the postmarginal vein is very short or absent. Macroteleia and Triteleia share the presence of a long marginal vein, though in the latter genus it is variable and the marginal and stigmal veins can be of similar length. However, both Macroteleia and Triteleia have a well-developed postmarginal vein. Habroteleia also differs from these genera in that it has a Ceratobaeus-type ovipositor (Austin & Field, 1997). The complexity of this system suggests that while these three genera appear to be quite similar, in fact they may not be closely related at all. Alternatively, it implies that the ovipositor system is much more labile than expected. Unfortunately, Habroteleia was not included among the taxa in the phylogenetic analysis of Murphy et al. (2007), and we therefore do not have an independent assessment of its relations. The structure of the ovipositor is of limited use for separating Habroteleia from Triteleia because it is rarely extruded in preserved specimens of the latter, and it is not obvious from external morphology (e.g. visibility of T7 in females) that Habroteleia has a Ceratobaeus-type ovipositor. Chen et al. (2013) provided a key to separate these genera, which we here present again.

Key to separate Macroteleia, Triteleia and Habroteleia 1 Postmarginal vein in fore wing absent or rudimentary; ovipositor Ceratobaeus-type ...... Habroteleia Kieffer - Postmarginal vein in fore wing well developed, distinctly longer than stigma vein (r-rs); ovipositor Scelio-type ...... 2 2 Female T6 strongly compressed laterally, wedge-like; male apical tergite apically emarginate or with 1 central spine but never bispinose...... Macroteleia Westwood - Female T6 triangular, not compressed laterally; male apical tergite with posterolateral conrners bispinose or at least pointed ...... Triteleiea Kieffer

Key to Females (unknown for H. ruficoxa (Kieffer)) 1 T1 with horn (Figs 2.3D, 2.4D, 2.5D, 2.6D, 2.7D, 2.11D, 2.13F) ...... 2

182 - T1 without horn (Figs 2.3F, 2.4F, 2.14C, 2.15F) ...... 4 2 Posterior vertex largely smooth with sparse to moderate punctures above occipital carina (Fig. 2.13D); gena sparsely punctate (Fig. 2.13B); mesepisternum anteroventral to mesopleural depression largely smooth with sparse punctures (Fig. 2.13B) ...... Habroteleia salebra Chen & Talamas, n. sp. - Posterior vertex densely punctate to punctate rugose (Figs 2.3C, 2.4D, 2.5D, 2.6D, 2.7D, 2.11C); gena densely punctate to punctate rugose (Figs 2.3B, 2.4B, 2.5B, 2.6B, 2.7B, 2.11B); mesepisternum anteroventral to mesopleural depression densely punctate to punctate rugose (Figs 2.3B, 2.4B, 2.5B, 2.6B, 2.7B, 2.11B) ...... 3 3 Median propodeal projection short (Figs 2.3D, 2.4D, 2.5D, 2.6D, 2.7D); T6 in female longitudinally striate, with fine punctures in interstices (Fig. 2.16A) ...... Habroteleia flavipes Kieffer - Median propodeal projection long (Figs 2.10D, 2.11D); T6 in female densely punctate and without longitudinal striae (Fig. 2.16B) ...... Habroteleia persimilis (Kozlov & Kononova) 4 Central keel of frons present (Figs 2.14D, 2.15E); upper frons densely punctate (Figs 2.14D, 2.15E); transverse sulcus on T2 present (Figs 2.14C, 2.15F) ...... 5 - Central keel of frons absent (Figs 2.8E, 2.9E); upper frons sparsely punctate (Figs 2.8E, 2.9E); transverse sulcus on T2 absent (Figs 2.3F, 2.4F) ...... Habroteleia mutabilis Chen& Talamas, n. sp. 5 Apex of T6 in female rounded (Fig. 2.14E); posterior vertex punctate rugose (Fig. 2.14D) ...... Habroteleia soa Chen & Talamas, n. sp. - Apex of T6 in female with small spine (Fig. 2.1E); posterior vertex smooth with sparse punctures (Fig. 2.15D) ...... Habroteleia spinosa Chen & Johnson, n. sp.

Key to Males 1 Apex of T8 with apical spine (Fig. 2.1C) ...... 2 - Apex of T8 without apical spine (Fig. 2.1D) ...... 4 2 Occipital carina interrupted medially (Fig. 2.13D); posterior vertex largely smooth with sparse to moderate punctures above occipital carina (Fig. 2.13D); gena sparsely punctate (Fig. 2.13B) ...... Habroteleia salebra Chen & Talamas, n. sp.

183 - Occipital carina complete (Figs 2.3D, 2.4D, 2.5D, 2.6D, 2.7D, 2.14D); posterior vertex densely punctate or punctate rugose (Figs 2.3D, 2.4D, 2.5D, 2.6D, 2.7D, 2.14C); gena densely punctate or punctate rugose (Figs. 2.3B, 2.5B, 2.6B, 2.7B, 2.14B) ...... 3 3 Central keel absent (Figs 2.4E, 2.5E); netrion rugulose anteriorly, smooth posteriorly, sometimes smooth only along posterior margin (Fig. 2.1F); T1 densely longitudinally striate with rugulose interstices (Figs 2.4F, 2.6F, 2.7F) ...... Habroteleia flavipes Kieffer - Central keel present (Fig. 2.14D); netrion coarsely striate (Fig. 2.14B); T1 sparsely longitudinally striate, smooth in interstices (Fig. 2.14C) ...... Habroteleia soa Chen & Talamas, n. sp. 4 Central keel absent (Figs 2.8E, 2.9E); transverse sulcus on T2 absent (Figs 2.8F, 2.9F) ...... Habroteleia mutabilis Chen & Talamas, n. sp. - Central keel present (Figs 2.10E, 2.12E, 2.15E); transverse sulcus on T2 present (Figs 2.11F, 2.12F, 2.15F) ...... 5 5 Median propodeal projection long (Figs 2.10D, 2.11B, 2.11D); notaulus formed by contiguous punctures (Figs 2.10D, 2.11C) .. Habroteleia persimilis (Kozlov & Kononova) - Median propodeal projection short (Figs 2.12B, 2.15D); notaulus formed by discrete punctures (Figs 2.12D, 2.15D) ...... 7 7 Posterior vertex punctate rugose (Fig. 2.12D); mesoscutal midlobe densely punctate (Fig. 2.12D); gena punctate rugose throughout (Fig. 2.12B) ...... Habroteleia ruficoxa (Kieffer) - Posterior vertex smooth with sparse punctures (Fig. 2.15D); mesoscutal midlobe densely and finely punctate along anterior margin, otherwise smooth (Fig. 2.15D): gena sparsely punctate (Fig. 2.15B) ...... Habroteleia spinosa Chen & Johnson, n. sp.

Species descriptions

Baryconus vindhiensis (Sharma), n. comb. http://zoobank.org/0EC31368-F49B-4183-B2C5-BD09C1C07753 http://bioguid.osu.edu/xbiod_concepts/4540 Figures 2.2A–2.2D

184 Triteleia vindhiensis Sharma, 1981: 451 (original description); Mani & Sharma, 1982: 168 (description, generic transfer). Habroteleia vindhiensis (Sharma): Johnson, 1992: 399 (cataloged, type information).

Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=4540] Material Examined. Holotype, female, T. vindhiensis: INDIA: Madhya Pradesh St., 21.5, Panna-Satna Road, 9.IX–10.IX.1979, M. S. Mani et al., USNMENT01197073 (deposited in USNM). Comments. The deep frontal depression margined by a sharp carina (Fig. 8), pronounced occiput (Fig. 9), long postmarginal vein and short marginal vein (Fig. 9) clearly indicate that this species belongs to Baryconus.

Habroteleia flavipes Kieffer http://zoobank.org/ACD49F55-9F4E-4C91-A044-22DCC0428FF6 http://bioguid.osu.edu/xbiod_concepts/4535 Figures 2.1F, 2.3A–2.7F, 2.16A

Habroteleia flavipes Kieffer, 1905: 15 (original description, keyed); Kieffer, 1926: 363 (description, keyed); Bin, 1974: 455 (type information); Johnson, 1992: 399 (cataloged, type information). Habroteleia browni Crawford, 1910: 125 (original description); Kieffer, 1926: 363, 364 (description, keyed); Baltazar, 1966: 177 (cataloged, synonymy, type information, distribution); Masner & Muesebeck, 1968: 37 (type information); Johnson, 1992: 399 (cataloged, type information), n. syn. http://zoobank.org/EC09DB18-92D9-4FB1-B986-3F7EAD7D54E4 http://bioguid.osu.edu/xbiod_concepts/4534 Chrestoteleia Bakeri Kieffer, 1913: 389 (original description); Kelner-Pillault, 1958: 150 (type information); Johnson, 1992: 399 (type information), n. syn. http://zoobank.org/F18A3905-9A5A-4755-A56A-5379E8564044 http://bioguid.osu.edu/xbiod_concepts/8935

185 Chrestoteleia bakeri Kieffer: Kieffer, 1926: 443 (description, keyed); Baltazar, 1966: 177 (junior synonym of Habroteleia browni Crawford); Baltazar, 1966: 182 (cataloged, type information, distribution). Habroteleia bakeri (Kieffer): Baltazar, 1961: 395 (generic transfer, diagnosis). Habroteleia bharatensis Saraswat, 1978: 7 (original description); Mani & Sharma, 1982: 167 (description); Johnson, 1992: 398 (cataloged), n. syn. http://zoobank.org/309A96B1-1DCA-45CA-B1AB-1D6E570C7E07 http://bioguid.osu.edu/xbiod_concepts/4533 Triteleia kotturensis Sharma, 1981: 447 (original description), n. syn. http://zoobank.org/28DFECE9-8723-4ACA-BB61-96B11C9546A8 http://bioguid.osu.edu/xbiod_concepts/8940 Habroteleia kotturensis (Sharma): Mani & Sharma, 1982: 168 (description, generic transfer); Johnson, 1992: 399 (cataloged, type information)

Description. Body length of female: 4.36–4.72 mm (n=20). Body length of male: 4.15–4.52 mm (n=20).Length of A3 in male: longer than A2. Punctation of frons above antennal scrobe: dense. Sculpture of antennal scrobe: punctate rugose to smooth. Central keel: absent. Sculpture of ventrolateral frons: punctate rugose. Occipital carina: complete. Sculpture of posterior vertex: densely punctate to punctate rugose. Sculpture of gena: densely punctate to punctate rugose. Sculpture of occiput: punctate rugose. Color of mesosoma: black. Sculpture of dorsal pronotal area: punctate rugose. Sculpture of lateral pronotal area: densely punctate. Sculpture of netrion: anterior half rugulose, posterior half smooth. Setae of netrion: dense throughout. Sculpture of notaulus: contiguously punctate. Sculpture of mesoscutal midlobe: largely punctate rugose, with a medial furrow and smooth areas laterally. Sculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of lateral propodeal area: rugose. Setation of mesoscutellum: dense. Sculpture of mesoscutellum: coarsely punctate rugose. Median propodeal projection: short. Mesopleural carina: distinct. Sculpture of mesepisternum anteroventral to mesopleural depression: punctate rugose. Sculpture of dorsal metapleural area: smooth to rugulose. Sculpture of ventral metapleural area: punctate rugose. Setation of ventral metapleural area: dense. Color of legs: orange-yellow to dark brown; dark brown to black. Sculpture of hind coxa: densely punctate.

186 Color of metasoma: black; black with T3–T4 and S2–S5 partly brown to yellow. T1 horn in female: present. Sculpture of posterior margin of T1 in female: densely longitudinally striate, punctate rugulose in interstices. Transverse sulcus on T2: present. Sculpture of T2–T5: densely longitudinally striate, with fine punctures in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: distinctly longer than wide. Apex of T6 in female: round. Sculpture of S2: longitudinally striate rugose. Sculpture of T1 in male: densely longitudinally striate, punctate rugulose in interstices. Male T8 apical spine: present. Diagnosis. This species is most similar to H. persimilis but can be distinguished by its short median propodeal projection and longitudinally striate T6 in female. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=4535] Material Examined. Holotype of Habroteleia flavipes Kieffer, female: INDONESIA: Sumatera Utara Prov., Sumatra Isl., Pangherang Pisang, X.1890 – III.1891, E. Modigliani, MCSN 0001 (deposited in MCSN). Holotype of Habroteleia bharatensis Saraswat, female: INDIA: West Bengal St., 16.4, Poro North, 6.IV–24.IV.1976, M. S. Mani, USNMENT01197132 (deposited in USNM). Syntype of Chrestoteleia bakeri Kieffer, female: PHILIPPINES: Laguna Prov., Los Baños, no date, Baker, ANIC DB 32-020728 (deposited in ANIC). Syntype of Chrestoteleia Bakeri Kieffer, female: PHILIPPINES: Laguna Prov., Los Baños, no date, Baker, MNHN 0013 (deposited in MNHN). Holotype of Habroteleia browni Crawford, female: PHILIPPINES: Metropolitan Manila Reg., Manila, no date, R. Brown, USNM Type No. 12894 (deposited in USNM). Holotype of Triteleia kotturensis Sharma, female: INDIA: Kerala St., 24.8, Kottur, 4.X.1980, M. S. Mani et al., USNMENT01197074 (deposited in USNM). Other material: (137 females, 79 males, 1 unknown) BANGLADESH: 2 females, OSUC 688056–688057 (CNCI). BRUNEI: 1 female, OSUC 232932 (BPBM). CAMBODIA: 1 female, OSUC 232935 (BPBM). CHINA: 23 females, 7 males, OSUC 232920 (BPBM); SCAU 2010100389, 2010100402, 2010100419, 2010100431, 2010100437, 2010100445–2010100446, 2010100459, 2010100464, 2010100495, 2010100497, 2010100499, 2010100502, 2010100504–2010100505, 2010100508– 2010100512, 2010100514, 2010100517–2010100518, 2010100521–2010100522, 2010100524– 2010100526, 2010100552 (SCAU). INDIA: 1 male, OSUC 688053 (CNCI). INDONESIA: 58 females, 23 males, OSUC 232906 (BPBM); OSUC 687960–688009, 688014–688041 (CNCI); OSUC 58007–58008 (OSUC). LAOS: 2 females, 3 males, OSUC 687955–687959 (CNCI).

187 MALAYSIA: 23 females, 26 males, OSUC 232907–232914, 232916–232919, 232923, 232931, 232933-232934, 232937, 246583 (BPBM); OSUC 687944–687954, 688058-688059 (CNCI); OSUC 491881–491896, 536427 (OSUC); OSUC 179084 (UCDC). PHILIPPINES: 1 female, 3 males, OSUC 232925–232928 (BPBM). SOUTH KOREA: 7 females, 2 males, 1 unknown, OSUC 687939 (CNCI); USNMENT01335741, 01335743–01335745, 01335747–01335749 (FSCA); USNMENT01335740, 01335742 (OSUC). SRI LANKA: 1 male, OSUC 688055 (CNCI). THAILAND: 16 females, 13 males, OSUC 232921–232922, 232924 (BPBM); OSUC 688042, 688049–688051 (CNCI); OSUC 321998–322002, 370249, 374199–374201, 381766– 381770, 688080–688087 (OSUC). VIETNAM: 3 females, OSUC 232915 (BPBM); OSUC 688052 (CNCI); OSUC 284756 (OSUC). Comments. The metasomal color in H. flavipes varies from entirely dark brown to having T3– T4 and S2–S5 mostly yellow, apparently without any correlation with geography. In males, the length of the spine at the apex of T8 varies from short to long, but it is always present.

Habroteleia impressa (Kieffer) http://zoobank.org/5A7AAB83-B2A4-401F-A137-D96D29D7648E http://bioguid.osu.edu/xbiod_concepts/4536

Crestoteleia impressa Kieffer, 1916: 180, 181 (original description, keyed, spelling error). Chrestoteleia impressa Kieffer: Kieffer, 1926: 443 (description, keyed); Baltazar, 1966: 182 (cataloged, distribution). Habroteleia impressa (Kieffer): Baltazar, 1966: 177 (cataloged, generic transfer, distribution); Johnson, 1992: 399 (cataloged, type information).

Comments. We were unable to locate the type specimens of this species, and its status and identity are unclear.

Habroteleia mutabilis Chen & Talamas, new species http://zoobank.org/5ADA1AD2-2B82-4314-A7A6-E65EBDBBE561 http://bioguid.osu.edu/xbiod_concepts/448460 Figures 2.1A–2.1B, 2.8A–2.9F

188

Description. Body length of female: 3.60–3.74 mm (n=20). Body length of male: 3.36–3.72 mm (n=20). Length of A3 in male: longer than A2.Punctation of frons above antennal scrobe: sparse. Sculpture of antennal scrobe: foveate. Central keel: absent. Sculpture of ventrolateral frons: densely punctate. Occipital carina: complete. Sculpture of posterior vertex: punctate rugose. Sculpture of gena: punctate rugose ventrally, sparsely punctate dorsally. Sculpture of occiput: smooth. Color of mesosoma: black; orange. Sculpture of dorsal pronotal area: punctate rugose. Sculpture of lateral pronotal area: smooth anteriorly, foveate posteriorly. Sculpture of netrion: coarsely striate. Setae of netrion: sparse throughout. Sculpture of notaulus: discretely punctate. Sculpture of mesoscutal midlobe: coarsely carinate with two rows of contiguous coarse punctures; largely smooth, with two rows of discrete coarse punctures. Sculpture of lateral lobe of mesoscutum: smooth. Sculpture of lateral propodeal area: foveate. Setation of mesoscutellum: sparse. Sculpture of mesoscutellum: sparsely punctate. Median propodeal projection: short; long. Mesopleural carina: distinct. Sculpture of mesepisternum anteroventral to mesopleural depression: smooth with a row of punctures along mesopleural carina. Sculpture of dorsal metapleural area: smooth. Sculpture of ventral metapleural area: smooth to foveate. Setation of ventral metapleural area: sparse. Color of legs: orange-yellow to dark brown; dark brown to black. Sculpture of hind coxa: smooth. Color of metasoma: black; orange with dark brown to black patches. T1 horn in female: absent. Sculpture of posterior margin of T1 in female: sparsely longitudinally striate. Transverse sulcus on T2: absent. Sculpture of T2–T5: T2–T3 sparsely longitudinally striate throughout, T4– T5 smooth medially, longitudinally striate. Sculpture of T6 in female: smooth. Length of T6 in female: wider than long. Apex of T6 in female: round. Sculpture of S2: sparsely longitudinally striate. Sculpture of T1 in male: sparsely longitudinally striate, smooth in interstices. Male T8 apical spine: absent. Etymology. The epithet is inspired by the Latin word for changeable, in reference to the variations in body color, sculpture of mesoscutal midlobe, and the length of median propodeal projection, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=448460]

189 Material Examined. Holotype, female: FIJI: Northern Div., Bua Prov., Vanua Levu Isl., 6km NW Kilaka Village, MT5, 98m, 16.807°S 178.991°E, Batiqere Range, 28.VI–21.VII.2004, Malaise trap, Schlinger & Tokota'a, FBA 142603 (deposited in BPBM). Paratypes: FIJI: 53 females, 27 males, FBA 070892, OSUC 232898, OSUC 232901, OSUC 232902, OSUC 232903, OSUC 232904, OSUC 232905 (BPBM); FBA 014394, 014404, 014409, 014413-014414, 019832, 025807, 025815, 029311, 029313, 029315, 029318-029320, 029323, 032077, 032086, 036322, 036328, 047849, 047855, 058998, 059005, 059026, 070887, 070893-070894, 082922, 084174-084175, 084181, 084183, 088442, 094483, 094485, 094487, 099217, 101111, 101129, 101568, 140945, 140956, 140961, 142806, 143124, 143130, 143134, 143209, 143217-143219, 144459, 151785, 151788, 164303, 166124, 166126, 166129, 166160, 166162-166163, 179833, 179838, 182136, 182139-182140, 182142, 186114, 188585, 188680, OSUC 688078, OSUC 688161, OSUC 688162 (CNCI). Other material: FIJI: 1 female, 1 male, OSUC 232900 (BPBM); FBA 084185 (CNCI). Comments. This species is well supported by many characters, although the color of mesosoma and metasoma, sculpture of mesoscutal midlobe, and the length of median propodeal projection are variable. The color of mesosoma and metasoma varies from orange to dark brown. The sculpture of the mesoscutal midlobe varies from largely smooth with two rows of discrete coarse punctures to coarsely carinate with two rows of contiguous coarse punctures. The length median propodeal projection varies from short to long. These variations are gradual among specimens. Therefore, we consider them as intraspecific rather than interspecific differences.

Habroteleia persimilis (Kozlov & Kononova) http://zoobank.org/F7A438F7-5207-4305-9467-11E23AA0923F http://bioguid.osu.edu/xbiod_concepts/243852 Figures 2.10A–2.11F, 2.16B–2.16C

Triteleia persimilis Kozlov & Kononova, 1985: 15, 17 (original description. Keyed); Kozlov & Kononova, 1990: 174, 178 (description, keyed); Johnson, 1992: 509 (cataloged, type information); Kononova, 1995: 69 (keyed); Kononova & Petrov, 2000: 28 (keyed). Habroteleia persimilis (Kozlov & Kononova): Kononova & Kozlov, 2008: 255 (description, generic transfer).

190 Triteleia dagavia Kozlov & Lê, 1995: 441, 445 (original description, keyed); Kozlov & Lê, 1996: 9, 14 (described as new, keyed); Lê, 2000: 76, 341 (description, keyed, type information), n. syn. http://zoobank.org/451262B6-B23F-487F-A870-AAB91CB1E35A http://bioguid.osu.edu/xbiod_concepts/28154 Habroteleia dagavia (Kozlov & Lê): Talamas & Pham, 2017: 227 (type information, generic transfer).

Description. Body length of female: 4.75–5.18 mm(n=20). Body length of male: 4.25–4.74 mm (n=20). Length of A3 in male: longer than A2. Punctation of frons above antennal scrobe: dense. Sculpture of antennal scrobe: punctate rugose to smooth. Central keel: present. Sculpture of ventrolateral frons: punctate rugose. Occipital carina: complete. Sculpture of posterior vertex: punctate rugose. Sculpture of gena: punctate rugose. Sculpture of occiput: densely finely punctate. Color of mesosoma: black. Sculpture of dorsal pronotal area: punctate rugose. Sculpture of lateral pronotal area: smooth anteriorly, foveate posteriorly. Sculpture of netrion: coarsely striate. Setae of netrion: sparse throughout. Sculpture of notaulus: contiguously punctate. Sculpture of mesoscutal midlobe: largely densely punctate, with a medial furrow and smooth areas laterally. Sculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of lateral propodeal area: rugose. Setation of mesoscutellum: dense. Sculpture of mesoscutellum: coarsely punctate rugose. Median propodeal projection: long. Mesopleural carina: distinct. Sculpture of mesepisternum anteroventral to mesopleural depression: punctate rugose. Sculpture of dorsal metapleural area: smooth. Sculpture of ventral metapleural area: rugose. Setation of ventral metapleural area: dense. Color of legs: orange-yellow. Sculpture of hind coxa: densely punctate. Color of metasoma: black. T1 horn in female: present. Sculpture of posterior margin of T1 in female: largely longitudinally striate with horn punctate rugose. Transverse sulcus on T2: present. Sculpture of T2–T5: densely longitudinally striate, with fine punctures in interstices. Sculpture of T6 in female: densely punctate. Length of T6 in female: distinctly longer than wide. Apex of T6 in female: round. Sculpture of S2: densely longitudinally striate, punctate in interstices. Sculpture of T1 in male: densely longitudinally striate, punctate rugulose in interstices. Male T8 apical spine: absent.

191 Diagnosis. This species is most similar to H. flavipes but can be distinguished by its long median propodeal projection and densely punctate T6 in female. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=243852] Material Examined. Holotype, female, T. persimilis: JAPAN: Aichi Pref., Honshu Isl., Inuyama City, 6.X.1981, E. Sugonyaev, ZIN 0014 (deposited in ZIN). Holotype of Triteleia dagavia Kozlov & Lê, female: VIETNAM: Quang Nam Prov., Lang Stream, forest, Dak Pring, 31.X.1979, X. H. Lê, IEBR 0143 (deposited in IEBR). Other material: (48 females, 43 males) CHINA: 6 females, 10 males, SCAU 2010100315–2010100317, 2010100319–2010100320, 2010100322, 2010100330, 2010100335, 2010100337–2010100340, 2010100347, 2010100349, 2010100352–2010100353 (SCAU). JAPAN: 40 females, 29 males, OSUC 687863, 687865– 687909, 687914–687936 (CNCI). SOUTH KOREA: 2 females, 4 males, OSUC 687937– 687938, 687940–687943 (CNCI). Comments. Habroteleia persimilis, like H. flavipes, has a distribution that spans a large latitudinal range, extending from central Vietnam into the Palearctic region in Japan and South Korea.

Habroteleia ruficoxa (Kieffer) http://zoobank.org/C3EF6C6D-486C-47FC-B481-81EDF21FA806 http://bioguid.osu.edu/xbiod_concepts/4538 Figures 2.12A–2.12F

Phaedroteleia ruficoxa Kieffer, 1916: 182, 183 (original description. Keyed); Kieffer, 1926: 418 (description, keyed); Kelner-Pillault, 1958: 151 (type information); Baltazar, 1966: 181 (cataloged, type information, distribution). Habroteleia ruficoxa (Kieffer): Masner, 1976: 26 (generic transfer); Johnson, 1992: 399 (cataloged, type information).

Description. Body length of male: 4.0 mm (n=1). Length of A3 in male: as long as A2. Punctation of frons above antennal scrobe: sparse. Sculpture of antennal scrobe: foveate. Central keel: present. Sculpture of ventrolateral frons: punctate rugose. Occipital carina: complete.

192 Sculpture of posterior vertex: punctate rugose. Sculpture of gena: punctate rugose. Sculpture of occiput: smooth. Color of mesosoma: black. Sculpture of dorsal pronotal area: punctate rugose. Sculpture of lateral pronotal area: smooth anteriorly, foveate posteriorly. Sculpture of netrion: coarsely striate. Setae of netrion: dense throughout. Sculpture of notaulus: discretely punctate. Sculpture of mesoscutal midlobe: densely punctate. Sculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of lateral propodeal area: rugose. Setation of mesoscutellum: sparse. Sculpture of mesoscutellum: sparsely punctate. Median propodeal projection: short. Mesopleural carina: distinct. Sculpture of mesepisternum anteroventral to mesopleural depression: largely smooth with sparse punctures. Sculpture of dorsal metapleural area: rugose. Sculpture of ventral metapleural area: rugose. Setation of ventral metapleural area: dense. Color of legs: orange- yellow. Sculpture of hind coxa: densely punctate. Color of metasoma: black. Transverse sulcus on T2: present. Sculpture of T2–T5: sparsely longitudinally striate, smooth in interstices. Sculpture of T1 in male: sparsely longitudinally striate, smooth in interstices. Male T8 apical spine: absent. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=4538] Material Examined. Holotype, male, P. ruficoxa: PHILIPPINES: Mindanao Isl., Butuan Chartered City, no date, Baker, MNHN_EY3427 (deposited in MNHN). Comments. The holotype specimen of Habroteleia ruficoxa is in reasonably good condition in that the characters used for diagnosis at the species level are readily accessible. The challenge is that the species was described from a single male and in the course of this revision we did not encounter any additional specimens of H. ruficoxa. The absence of a spine on T8 in the male, the largely smooth surface of the mesoscutum and mesoscutellum, and the notauli weakly indicated by punctures place the holotype specimen well outside of our concept of H. flavipes, the only other species of Habroteleia known from the Philippines.

Habroteleia salebra Chen & Talamas, new species http://zoobank.org/F60BFB76-6AA1-4484-B2C7-CA6BE93CED9F http://bioguid.osu.edu/xbiod_concepts/448456 Figures 2.1C, 2.13A–2.13F

193 Description. Body length of female: 4.28–4.90 mm (n=20). Body length of male: 4.30–4.73mm (n=20). Length of A3 in male: longer than A2. Punctation of frons above antennal scrobe: sparse. Sculpture of antennal scrobe: smooth. Central keel: absent. Sculpture of ventrolateral frons: punctate rugose. Occipital carina: interrupted medially. Sculpture of posterior vertex: smooth with sparse punctures. Sculpture of gena: sparsely punctate. Sculpture of occiput: densely finely punctate. Color of mesosoma: black. Sculpture of dorsal pronotal area: punctate rugose. Sculpture of lateral pronotal area: smooth anteriorly, foveate posteriorly. Sculpture of netrion: coarsely striate. Setae of netrion: sparse throughout. Sculpture of notaulus: discretely punctate. Sculpture of mesoscutal midlobe: densely and finely punctate along anterior margin, sparsely punctate along posterior margin, otherwise smooth. Sculpture of lateral lobe of mesoscutum: smooth. Sculpture of lateral propodeal area: rugose. Setation of mesoscutellum: sparse. Sculpture of mesoscutellum: coarsely punctate rugose. Median propodeal projection: short. Mesopleural carina: weakly developed. Sculpture of mesepisternum anteroventral to mesopleural depression: largely smooth with sparse punctures. Sculpture of dorsal metapleural area: smooth. Sculpture of ventral metapleural area: punctate rugose. Setation of ventral metapleural area: dense. Color of legs: dark brown to black. Sculpture of hind coxa: densely punctate. Color of metasoma: black. T1 horn in female: present. Sculpture of posterior margin of T1 in female: densely longitudinally striate, punctate rugulose in interstices. Transverse sulcus on T2: present. Sculpture of T2–T5: densely longitudinally striate, with fine punctures in interstices. Sculpture of T6 in female: densely punctate. Length of T6 in female: distinctly longer than wide. Apex of T6 in female: round. Sculpture of S2: sparsely longitudinally striate medially, with fine punctures in interstices, irregularly finely punctate laterally. Sculpture of T1 in male: densely longitudinally striate, punctate rugulose in interstices. Male T8 apical spine: present. Diagnosis. This species is most similar to H. spinosa but can be distinguished by the round apex of T6 in females and the absence of a spine on the apex of T8 in males. Etymology. The epithet is inspired by the Latin word for a rough, uneven road, in reference to the glabrous netrion sulcus adjacent to the setose posterior portion of the netrion, and is intended to be treated as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=448456]

194 Material Examined. Holotype, female: PAPUA NEW GUINEA: Madang Prov., 100m, 04°16'S 144°58'E, Morox, 1.VIII–18.VIII.2006, yellow pan trap, V. Iwam, OSUC 688076 (deposited in CNCI). Paratypes: (21 females, 12 males) INDONESIA: 1 female, OSUC 232875 (BPBM). PAPUA NEW GUINEA: 20 females, 12 males, OSUC 232876–232877, 232879– 232884, 232886, 232890–232892, 232894–232897 (BPBM); OSUC 688060–688063, 688065– 688075, 688077 (CNCI).

Habroteleia scapularis (Kieffer) http://zoobank.org/6008D3A5-FE3A-4C26-8E5A-455A00D5DB9A http://bioguid.osu.edu/xbiod_concepts/4539

Crestoteleia scapularis Kieffer, 1916: 180 (original description, keyed, spelling error). Chrestoteleia scapularis Kieffer: Kieffer, 1926: 443, 444 (description, keyed); Baltazar, 1966: 182 (cataloged, distribution). Habroteleia scapularis (Kieffer): Baltazar, 1966: 177 (cataloged, generic transfer, distribution); Johnson, 1992: 399 (cataloged, type information).

Comments. We were not able to locate the type specimens of this species, and its status and identity are unclear.

Habroteleia soa Chen & Talamas, new species http://zoobank.org/DD68E31A-9B97-4226-832E-2549DD5F0E0A http://bioguid.osu.edu/xbiod_concepts/448556 Figures 2.14A–2.14E

Description. Body length of female: 3.72 mm (n=1). Length of A3 in male: longer than A2.Length of A3 in male: longer than A2. Punctation of frons above antennal scrobe: dense. Sculpture of antennal scrobe: smooth. Central keel: present. Sculpture of ventrolateral frons: punctate rugose. Occipital carina: complete. Sculpture of posterior vertex: punctate rugose. Sculpture of gena: punctate rugose ventrally, sparsely punctate dorsally. Sculpture of occiput: rugulose.

195 Color of mesosoma: black. Sculpture of dorsal pronotal area: punctate rugose. Sculpture of lateral pronotal area: smooth anteriorly, foveate posteriorly. Sculpture of netrion: coarsely striate. Setae of netrion: sparse throughout. Sculpture of notaulus: contiguously punctate. Sculpture of mesoscutal midlobe: punctate rugose on the anterior margin, otherwise largely smooth with two rows of discrete punctures. Sculpture of lateral lobe of mesoscutum: smooth. Sculpture of lateral propodeal area: rugose. Setation of mesoscutellum: sparse. Sculpture of mesoscutellum: coarsely punctate rugose. Median propodeal projection: short. Mesopleural carina: weakly developed. Sculpture of mesepisternum anteroventral to mesopleural depression: smooth with a row of punctures along mesopleural carina. Sculpture of dorsal metapleural area: smooth. Sculpture of ventral metapleural area: punctate rugose. Setation of ventral metapleural area: sparse. Color of legs: orange-yellow to dark brown. Sculpture of hind coxa: smooth. Color of metasoma: black. T1 horn in female: absent. Sculpture of posterior margin of T1 in female: sparsely longitudinally striate. Transverse sulcus on T2: present. Sculpture of T2–T5: T2–T4 sparsely longitudinally striate, with fine punctures in interstices, T5 densely longitudinally striate punctate. Sculpture of T6 in female: densely punctate. Length of T6 in female: wider than long. Apex of T6 in female: round. Sculpture of S2: longitudinally striate rugose. Sculpture of T1 in male: sparsely longitudinally striate, smooth in interstices. Male T8 apical spine: present. Etymology. The Malagasy word “soa” means “beautiful” or “excellent”. We apply it to this species because we find it to be both of these. The name is treated as a noun in apposition. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=448556] Material Examined. Holotype, female: MADAGASCAR: Antsiranana Auto. Prov., 5km W Manantenina, Camp Mantella, low altitude rainforest, MA-31-32, 490m, 14°26.29'S 49°46.44'E, Marojejy National Park, 14.X–22.X.2005, Malaise trap, M. Irwin & R. Harin'Hala, CASENT 2136859 (deposited in CSCA). Paratypes: MADAGASCAR: 3 males, CASENT 2132434– 2132435 (OSUC), 2135976 (CSCA). Comments. Habroteleia soa is the most geographically disjunct member of the genus, separated from the other species by the Indian Ocean. Despite this separation, it is not morphologically unusual in comparison with the other species, suggesting either that there is a relatively recent division between H. soa and the other species, that the morphology of the genus evolves rather

196 slowly, or that there has been insufficient sampling in the intervening areas (e.g., east Africa, the moist southern part of the Arabian peninsula, India, and all other intervening regions).

Habroteleia spinosa Chen & Johnson, new species http://zoobank.org/9DD4E72F-B7E1-42CE-95BF-DDA22297830C http://bioguid.osu.edu/xbiod_concepts/448458 Figures 2.1D–2.1E, 2.15A–2.15F

Description. Body length of female: 3.51–3.52 mm (n=2). Body length of male: 3.37–3.81mm (n=6).Length of A3 in male: longer than A2. Punctation of frons above antennal scrobe: dense. Sculpture of antennal scrobe: punctate rugose. Central keel: present. Sculpture of ventrolateral frons: punctate rugose. Occipital carina: interrupted medially. Sculpture of posterior vertex: smooth with sparse punctures. Sculpture of gena: sparsely punctate. Sculpture of occiput: smooth. Color of mesosoma: black. Sculpture of dorsal pronotal area: sparsely punctate. Sculpture of lateral pronotal area: largely smooth, with sparsely punctures medially. Sculpture of netrion: coarsely striate ventrally, rugulose dorsally. Setae of netrion: dense throughout. Sculpture of notaulus: discretely punctate. Sculpture of mesoscutal midlobe: densely finely punctate along anterior margin, otherwise smooth. Sculpture of lateral lobe of mesoscutum: smooth. Sculpture of lateral propodeal area: rugose. Setation of mesoscutellum: sparse. Sculpture of mesoscutellum: sparsely punctate. Median propodeal projection: short. Mesopleural carina: distinct. Sculpture of mesepisternum anteroventral to mesopleural depression: smooth. Sculpture of dorsal metapleural area: smooth. Sculpture of ventral metapleural area: rugose. Setation of ventral metapleural area: dense. Color of legs: dark brown to black. Sculpture of hind coxa: smooth. Color of metasoma: black. T1 horn in female: absent. Sculpture of posterior margin of T1 in female: densely longitudinally striate, punctate rugulose in interstices. Transverse sulcus on T2: present. Sculpture of T2–T5: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: rugose. Length of T6 in female: wider than long. Apex of T6 in female: pointed. Sculpture of S2: sparsely longitudinally striate medially, with fine punctures in

197 interstices, irregularly finely punctate laterally. Sculpture of T1 in male: sparsely longitudinally striate, smooth in interstices. Male T8 apical spine: absent. Diagnosis. This species is most similar to H. salebra but can be distinguished by the pointed apex of T6 in females and the spine at the apex of T8 in males, Etymology. The specific epithet means spiny, referring to the pointed apex of T6 in females and should be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=448458] Material Examined. Holotype, female: INDONESIA: Papua Prov., W New Guinea Isl., Central Mts., Archbold Lake, 760m, 26.XI–3.XII.1961, sweeping, L. W. Quate, OSUC 232889 (deposited in BPBM). Paratypes: (1 female, 6 males) INDONESIA: 6 males, OSUC 232878, 232887, 232893, 232929–232930, 234491 (BPBM). PAPUA NEW GUINEA: 1 female, OSUC 232888 (BPBM).

198

Figure 2.1 A–B, Habroteleia mutabilis n. sp. A, Paratype (FBA 143219), Propodeum, dorsolateral view; B, Holotype (FBA 070892), Propodeum, dorsolateral view; C, Habroteleia salebra n. sp., male, paratype (OSUC 688063), Apex of metasoma, dorsal view; D, Habroteleia spinosa n. sp., male, paratype (OSUC 232878), Apex of metasoma, dorsal view; E, Habroteleia spinosa n. sp., female, holotype (OSUC 232889), Apex of metasoma, dorsal view; F, Habroteleia flavipes, male (OSUC 58007), Pronotum, lateral view.

199

Figure 2.2 Baryconus vindhiensis, female, holotype (USNMENT01197073). A, Lateral habitus; B, Head, lateral view; C, Dorsal habitus; D, Labels.

200

Figure 2.3 Habroteleia flavipes, female, holotype (MCSN 0001). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, lateral view; F, Metasoma and wings, dorsal view.

201

Figure 2.4 Habroteleia bharatensis, female, holotype (USNMENT 01197132). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

202

Figure 2.5 Habroteleia browni, female, holotype (USNM Type No. 12894). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view.

203

Figure 2.6 Chrestoteleia bakeri, female, holotype (MNHN 0013). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view.

204

Figure 2.7 Habroteleia kotturensis, female, holotype (USNMENT01197074). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, lateral view; F, Metasoma and wings, dorsal view.

205

Figure 2.8 Habroteleia mutabilis n. sp., female, holotype (FBA 142603). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view.

206

Figure 2.9 Habroteleia mutabilis n. sp., female, paratype (FBA 070892). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view.

207

Figure 2.10 Triteleia persimilis, female, holotype (ZMAS 0139). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

208

Figure 2.11 Triteleia dagavia, female, holotype (IEBR 0143). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Head and mesosoma, dorsal view; D, Propodeum, dorsolateral view; E, Head, lateral view; F, Metasoma, dorsal view.

209

Figure 2.12 Phaedroteleia ruficoxa, male, holotype (MNHN_EY3427). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

210

Figure 2.13 Habroteleia salebra n. sp., female, holotype (OSUC 688076). A, Lateral habitus; B, Head and mesosoma, lateral view C, Dorsal habitus D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view.

211

Figure 2.14 Habroteleia soa n. sp., female, holotype (CASENT 2136859). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head, anterior view; E, T5 and T6, dorsal view.

212

Figure 2.15 Habroteleia spinosa n. sp., female, holotype (OSUC 232889). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma and wings, dorsal view.

213

Figure 2.16 A, Habroteleia flavipes, female (OSUC 688019), T5 and T6, dorsal view. B–C, Habroteleia persimilis, female (OSUC 687934). B, T5 and T6, dorsal view; C, Wings.

214 Table 2.1 List of morphological terms and links to Hymenoptera Anatomy Ontology database for world species of Habroteleia.

A1 http://purl.obolibrary.org/obo/HAO_0000908 A2 http://purl.obolibrary.org/obo/HAO_0000706 A3 http://purl.obolibrary.org/obo/HAO_0001148 A7 http://purl.obolibrary.org/obo/HAO_0001885 A12 http://purl.obolibrary.org/obo/HAO_0001884 antenna http://purl.obolibrary.org/obo/HAO_0000101 antennomere http://purl.obolibrary.org/obo/HAO_0000107 area http://purl.obolibrary.org/obo/HAO_0000146 body http://purl.obolibrary.org/obo/HAO_0000182 carina http://purl.obolibrary.org/obo/HAO_0000188 central keel http://purl.obolibrary.org/obo/HAO_0000109 cpa cervical pronotal area http://purl.obolibrary.org/obo/HAO_0000194 clava http://purl.obolibrary.org/obo/HAO_0000203 clypeus http://purl.obolibrary.org/obo/HAO_0000212 compound eye http://purl.obolibrary.org/obo/HAO_0000217 coxa http://purl.obolibrary.org/obo/HAO_0000228 depression http://purl.obolibrary.org/obo/HAO_0000241 dpa dorsal pronotal area http://purl.obolibrary.org/obo/HAO_0000267 egg http://purl.obolibrary.org/obo/HAO_0000286 epomial carina http://purl.obolibrary.org/obo/HAO_0000307 eye http://purl.obolibrary.org/obo/HAO_0000217 femur http://purl.obolibrary.org/obo/HAO_0000327 fore wing http://purl.obolibrary.org/obo/HAO_0000351 frons http://purl.obolibrary.org/obo/HAO_0001523 gena http://purl.obolibrary.org/obo/HAO_0000371 head http://purl.obolibrary.org/obo/HAO_0000397 hind coxa http://purl.obolibrary.org/obo/HAO_0000587 hind tibia http://purl.obolibrary.org/obo/HAO_0000631 hind wing http://purl.obolibrary.org/obo/HAO_0000400 inner orbit http://purl.obolibrary.org/obo/HAO_0000419 interantennal process http://purl.obolibrary.org/obo/HAO_0000422 lateral lobe of mesoscutum http://purl.obolibrary.org/obo/HAO_0000466 lateral ocellus http://purl.obolibrary.org/obo/HAO_0000481 LOL lateral ocellar line http://purl.obolibrary.org/obo/HAO_0000480 lpa lateral pronotal area http://purl.obolibrary.org/obo/HAO_0000483 malar sulcus http://purl.obolibrary.org/obo/HAO_0000504 mandible http://purl.obolibrary.org/obo/HAO_0000506 margin http://purl.obolibrary.org/obo/HAO_0000510 mesepisternum http://purl.obolibrary.org/obo/HAO_0001872 med mesopleural depression http://purl.obolibrary.org/obo/HAO_0000326 mesopleuron http://purl.obolibrary.org/obo/HAO_0000566

215 Table 2.1. Continued mesoscutellum http://purl.obolibrary.org/obo/HAO_0000574 mesoscutum http://purl.obolibrary.org/obo/HAO_0001490 mesosoma http://purl.obolibrary.org/obo/HAO_0000576 metapleuron http://purl.obolibrary.org/obo/HAO_0000621 metascutellum http://purl.obolibrary.org/obo/HAO_0000625 metasoma http://purl.obolibrary.org/obo/HAO_0000626 midlobe of mesoscutum http://purl.obolibrary.org/obo/HAO_0000520 netrion http://purl.obolibrary.org/obo/HAO_0000644 notauli (notaulus) http://purl.obolibrary.org/obo/HAO_0000647 occipital carina http://purl.obolibrary.org/obo/HAO_0000653 ocellus http://purl.obolibrary.org/obo/HAO_0000661 ot ocellar triangle http://purl.obolibrary.org/obo/HAO_0000430 OOL ocular ocellar line http://purl.obolibrary.org/obo/HAO_0000662 orbit http://purl.obolibrary.org/obo/HAO_0000672 POL posterior ocellar line http://purl.obolibrary.org/obo/HAO_0000759 process http://purl.obolibrary.org/obo/HAO_0000822 propodeum http://purl.obolibrary.org/obo/HAO_0001248 S1 http://purl.obolibrary.org/obo/HAO_0001997 S2 http://purl.obolibrary.org/obo/HAO_0001829 S3 http://purl.obolibrary.org/obo/HAO_0001831 S7 http://purl.obolibrary.org/obo/HAO_0002185 sculpture http://purl.obolibrary.org/obo/HAO_0000913 sternite http://purl.obolibrary.org/obo/HAO_0001654 sulcus http://purl.obolibrary.org/obo/HAO_0000978 T1 http://purl.obolibrary.org/obo/HAO_0000053 T2 http://purl.obolibrary.org/obo/HAO_0000056 T3 http://purl.obolibrary.org/obo/HAO_0000057 T4 http://purl.obolibrary.org/obo/HAO_0000058 T5 http://purl.obolibrary.org/obo/HAO_0000059 T6 http://purl.obolibrary.org/obo/HAO_0000060 T7 http://purl.obolibrary.org/obo/HAO_0000061 tergite http://purl.obolibrary.org/obo/HAO_0001783 tibia http://purl.obolibrary.org/obo/HAO_0001017 tyloid http://purl.obolibrary.org/obo/HAO_0001199 vein http://purl.obolibrary.org/obo/HAO_0001095 vertex http://purl.obolibrary.org/obo/HAO_0001077 vertical epomial carina http://purl.obolibrary.org/obo/HAO_0000307 wing http://purl.obolibrary.org/obo/HAO_0001089

216 Acknowledgments Thanks to L. Musetti and S. Hemly (OSUC) for essential assistance with specimen handling, software, and databasing; Agnièle Touret-Alby (MNHN) and Konstantin Samartsev (ZIN) for imaging type material. We thank the Florida Department of Agriculture and Consumer Services – Division of Plant Industry for their support on this contribution. This material is based upon work supported in part by the National Science Foundation of USA under grant No. DEB- 0614764 to N.F. Johnson and A.D. Austin.

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219 Chapter 3: Revision of the World species of the Genus Chromoteleia Ashmead (Hymenoptera, Platygastridae, Scelioninae)

Abstract The genus Chromoteleia Ashmead is revised. Twenty-seven species are recognized, six species are redescribed: C. congoana (Risbec), C. connectens Kieffer, C. fuscicornis Kieffer, C. longitarsis Kieffer, C. semicyanea Ashmead, C. tricarinata Kieffer; and twenty-one species are described as new: C. aequalis Chen & Johnson, sp. n., C. alternata Chen & Johnson, sp. n., C. bidens Chen & Masner, sp. n., C. copiosa Chen & Johnson, sp. n., C. cuneus Chen & Johnson, sp. n., C. curta Chen & Johnson, sp. n., C. depilis Chen & Johnson, sp. n., C. dispar Chen & Masner, sp. n., C. feng Chen & Johnson, sp. n., C. fossa Chen & Johnson, sp. n., C. ingens Chen & Masner, sp. n., C. levitas Chen & Johnson, sp. n., C. longa Chen & Johnson, sp. n., C. maura Chen & Masner, sp. n., C. parvitas Chen & Johnson, sp. n., C. pilus Chen & Johnson, sp. n., C. plana Chen & Johnson, sp. n., C. rara Chen & Johnson, sp. n., C. robusta Chen & Johnson, sp. n., C. semilutea Chen & Johnson, sp. n., C. sparsa Chen & Johnson, sp. n. Chromoteleia rufithorax Kieffer remains valid species but its identity and status are unclear. All species are known only from the Neotropical region except for Chromoteleia congoana (Resbec), which only occurs in Africa.

Introduction The genus Chromoteleia was originally described by Ashmead (1893) based on the colorful type species Chromoteleia semicyanea Ashmead collected in Saint Vincent. Later, Kieffer (1926) proposed Petalosema for species with expanded metanotum (metascutellum). Masner (1976) synonymized Petalosema with Chromoteleia by pointing out that Kieffer (1926) was misled by Ashmead’s (1893) inaccurate illustration of C. semicyanea showing a narrow and unexpanded metanotum. Six species were recorded from the New World: C. connectens Kieffer, C. fuscicornis Kieffer, C. longitarsis Kieffer, C. rufithorax Kieffer, C. semicyanea Ashmead, and C. tricarinata Kieffer. One species also was described from the Afrotropical region, C. congolana (Risbec), from what is currently Gabon. One fossil species, C. theobaldi Maneval, is reported from Baltic amber (from 40~60 mya) (Maneval 1938), but the status of this species is not clear (Talamas and Buffington 2015) and requires direct examination.

220 Since its original description in 1893, Chromoteleia has never been comprehensively reviewed. Although there are no host records for Chromoteleia, we suspect that it parasitizes the eggs of orthoptera based on large size and elongate shape of the parasitoids. A number of scelionine genera of similar habitus, and presumed close relation, are also known to be parasitoids of orthopteran eggs (e.g. Macroteleia Kieffer (Muesebeck, 1977), Triteleia Kieffer (Popovici et al. 2011)). The goal of this work is to produce a systematic revision of the world species of Chromoteleia, expand the biogeographic data associated with these species, and to present a clarified generic concept. The contributions of the authors are as follows. H.-y. Chen, E. J. Talamas, A. A. Valerio and N.F. Johnson: character definition, generic concept development, species concept development, imaging, key development, manuscript preparation; L. Masner: character definition, generic concept development, species concept development. The authors of the new species are indicated in the heading of each description.

Materials and methods

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; BPBM, Bernice P. Bishop Museum, Honolulu, HI; CAS, California Academy of Sciences, San Francisco, CA; CNCI, Canadian National Collection of Insects, Ottawa, Canada; CSCA, California State Collection of Arthropods, Sacramento, CA; FSCA, Florida State Collection of Arthropods, Gainesville, FL; IAVH, Instituto Alexander von Humboldt, Colección de Artrópodos, Villa de Leyva, Colombia; INHS, Illinois Natural History Survey, Urbana, IL; KUNH, Illinois Natural History Museum, Lawrence, KS; MIZA, Museo del Instituto de Zoología Agrícola, Maracay, Venezuela; MNHN, Muséum National d'Histoire Naturelle, Paris, France; MPEG, Museu Paraense Emílio Goeldi, Belém, PA, Brazil; MZLU, Lund Museum of Zoology, Lund University, Lund, Sweden; OSUC, C.A. Triplehorn Insect Collection, Ohio State University, Columbus, OH; SAMC, South African Museum, Iziko Museums of Cape Town, South Africa; TAMU, Texas A&M University Insect Collection, College Station, TX; UCDC, R.M. Bohart Museum of Entomology, University of California,

221 Davis, CA; UCMC, University of Colorado, Boulder, CO; UCRC, University of California, Riverside, CA; USNM, National Museum of Natural History, Washington, DC. 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); EH: eye height, length of compound eye measured parallel to dorsoventral midline of head; IOS: interocular space, minimal distance on frons between compound eyes; OD: ocellar diameter, greatest width of ocellus; OOL: ocular ocellar line, shortest distance from inner orbit and outer margin of posterior ocellus (Masner 1980); T1, T2, ... T7: metasomal tergite 1, 2, ... 7; S1, S2, … S7: metasomal sternite 1, 2, … 7. Morphological terminology otherwise generally follows Masner (1980) and Mikó et al. (2007). Morphological terms used in this work were as in the Hymenoptera Anatomy Ontology (Yoder et al. 2010) (Table 3.1). Identifiers (URIs) in the format HAO_XXXXXXX represent concepts in the HAO and are provided to enable readers to confirm their understanding of the concepts being referenced. To find out more about a given concept, including additional images, notes, references and other metadata, use the identifier as a search term at http://glossary.hymao.org or use the identifier as a web-link. In the Material Examined section the specimens studied are recorded in an abbreviated format, using unique identifiers (numbers prefixed with “OSUC”, “CASENT”, “FBA”, “MNHN_EY”) 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 can be accessed at the following link, hol.osu.edu, and entering the identifier in the form (note the space between the acronym and the number). 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, www.zoobank.org), and other taxonomic names, where appropriate, have been retrospectively registered. The external hyperlinks are explicitly cited in the endnotes so that users of the printed version of this article have access to the same resources.

222 Data associated with the genus Chromoteleia can be accessed at http://hol.osu.edu/index.html?id=464. The generic and species descriptions were generated by a database application, vSysLab (vsyslab.osu.edu), designed to facilitate the production of a taxon by character data matrices, and to integrate those data 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 are indicated by semicolon-separated character states. Images and measurements were made using Combine ZP and AutoMontage extended-focus software, using JVC KY-F75U digital camera, Leica Z16 APOA microscope, and 1X objective lens. Images were post-processed with Adobe Photoshop CS3 Extended. A standard set of images is provided for each species: dorsal habitus, lateral habitus, dorsal and lateral views of the head and mesosoma, and anterior view of head. The individual images are archived in Specimage (specimage.osu.edu), the image database at The Ohio State University.

Taxonomy

Chromoteleia Ashmead http://zoobank.org/25D9A544-B778-4365-9542-1E942D25AAF9 http://bioguid.osu.edu/xbiod_concepts/464

Chromoteleia Ashmead, 1893: 209, 211, 219Type: Chromoteleia semicyanea Ashmead, by monotypy and original designation (keyed); Ashmead, 1894: 216 (keyed); Dalla Torre, 1898: 501 (catalog of species); Ashmead, 1900: 327 (list of species of West Indies); Ashmead, 1903: 91, 93 (keyed); Kieffer, 1907: 266 (key to species); Brues, 1908: 26, 27, 28, 35 (diagnosis, keyed, list of species); Kieffer, 1908b: 115 (keyed); Kieffer, 1910a: 312 (key to species); Kieffer, 1910b: 62, 68 (description, list of species, keyed); Kieffer, 1913: 224 (description); Kieffer, 1926: 269, 406 (description, keyed); Muesebeck & Walkley, 1956: 342 (citation of type species); Masner, 1976: 23, 24 (description, synonymy; key to separate Baryconus Forster, Bracalba Dodd, Chromoteleia Ashmead, Oxyscelio Kieffer); Carpenter, 1992: 471 (fossil references); Johnson, 1992: 363 (cataloged, catalog of world species); Loiácono & Margaría, 2002: 557 (catalog of Brazilian species).

223 Petalosema Kieffer, 1926: 267, 358. Type: Chromoteleia rufithorax Kieffer, by original description (keyed, key to species), designated by Muesebeck & Walkley, 1956. Synonymized by Masner (1976); Muesebeck & Walkley, 1956: 336 (citation of type species); De Santis, 1980: 310: (catalog of species of Brazil). http://zoobank.org/E5906ABF-3A4D-4005-BFEC-13B2AEBD6E81 http://bioguid.osu.edu/xbiod_concepts/8521

Description. Length 3.38–9.20 mm; body elongate, robust. Head. Head shape in dorsal view: transverse. Vertex: densely punctate to punctate rugose. Hyperoccipital carina: absent. Occipital carina: present, complete or broadly interrupted medially. OOL: lateral ocellus nearly contiguous with inner orbits, OOL < 0.5 OD; lateral ocellus contiguous with inner orbit. Upper frons: convex, without frontal shelf or carina, punctate rugose. Antennal scrobe: broadly convex to concave medially with distinct depression. Submedian carina: absent. Orbital carina: absent. Inner orbits: diverging ventrally. IOS/EH: IOS distinctly less than EH. Interantennal process: short, often excavate medially. Central keel: present or absent. Antennal foramen: oriented laterally on interantennal process. Facial striae: absent. Malar sulcus: present. Malar striae: absent. Setation of compound eye: absent. Gena: broad, convex, distinctly produced behind eye. Clypeus shape: narrow, slightly convex medially, lateral corners not produced. Anterior (or ventral) margin of clypeus: pointed; straight. Labrum: not visible in anterior view. Number of mandibular teeth: 3. Arrangement of mandibular teeth: transverse. Number of maxillary palpomeres: 4. Shape of maxillary palpomeres: cylindrical. Number of labial palpomeres: 2. Antenna. 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 antennomeres with basiconic sensilla in female: 5; 6. Number of antennomeres with basiconic sensilla in female: 5; 6.Arrangement of sensilla on female clava: in longitudinal pairs. Number of antennomeres bearing tyloids in male antenna: 1. Shape of male flagellum: filiform. Mesosoma. Posterior apex of pronotum in dorsal view: straight, bifid apically to articulate with tegula. Epomial carina: present. Anterior face of pronotum: oblique, visible dorsally, short.

224 Lateral face of pronotum: weakly concave below position of dorsal epomial carina. Netrion: present. Netrion shape: moderately wide, open ventrally. Anterior portion of mesoscutum: vertical, flexed ventrally to meet pronotum. Mesoscutum shape: pentagonal, excavate at base of wings. Skaphion: absent. Notauli: present, percurrent. Parapsidal lines: absent. Antero-admedian lines: absent. Transscutal articulation: well-developed, narrow. Shape of mesoscutellum: trapezoidal, without spines. Lateral mesoscutellar spine: absent. Median mesoscutellar spine: absent. Axillular spine: absent. Surface of mesoscutellum: convex throughout. Median longitudinal furrow on mesoscutellum: absent. Metascutellum: clearly differentiated. Shape of metascutellum: trapezoidal with broad posterior margin; elongate trapezoidal but with deeply incised apex, forming two spines laterally. Posterior margin of metascutellum: straight; concave; convex. Setation of metascutellum: present. Metapostnotum: fused to propodeum. Lateral propodeal projection: absent. Medial propodeal projection: absent. Mesopleural carina: present. Mesal course of acetabular carina: not separating fore coxae. Setation of subalar pit: present. Mesopleural pit: present. Posterodorsal corner of mesopleuron: rounded anteriorly. Legs. Number of mesotibial spurs: 1. Number of metatibial spurs: 1. Dorsal surface of metacoxa: smooth; punctate. Shape of metacoxa: cylindrical, ecarinate. Trochantellus: indicated by transverse sulcus on femur. Wings. Wing development of female: macropterous. Wing development of male: macropterous. Tubular veins in fore wing: present. Bulla of fore wing R: absent. Length of marginal vein of fore wing: punctiform, R terminating at costal margin. Origin of r-rs in fore wing: basal of point at which R meets costal margin. Basal vein (Rs+M) in fore wing: spectral; nebulous. Development of R vein in hind wing: complete. Metasoma. Number of external metasomal tergites in female: 6. Number of external metasoma sternites in female: 6. Number of external metasomal tergites in male: 7. Number of external metasomal sternites in male: 7. Shape of metasoma: lanceolate. Laterotergites: present, narrow. Laterosternites: present. T1 of female: flat; medially convex as a small hump anteriorly. Relative size of metasomal segments: T2–T3 subequal in length, remaining terga shorter. Metasomal tergites with basal crenulae: T2. Sublateral carinae on tergites: present. Median longitudinal carina on metasomal terga: absent. Shape of female T6: flattened; laterally compressed. Anterior margin of S1: not produced anteriorly, straight. Felt fields on S2: absent. Felt fields on S3: absent. Ovipositor: Scelio-type (Austin & Field 1997).

225

Generic Diagnosis The large size and distinctive characters (metascutellum large and setose; propodeum without projections; marginal vein of forwing punctiform) of Chromoteleia make it a relatively easy genus to identify. The setation of the metascutellum is found in relatively few scelionine genera, particularly among the more robust genera, and is thus a very useful diagnostic character. Chromoteleia appears closest to Bracalba Dodd and Romilius Walker, from which it can be separated by the setation of the eyes (absent in Chromoteleia).

Comments: The distribution of Chromoteleia in Africa and South America is a phenomenon of biogeographical interest. Dispersal from South America to Africa has been demonstrated in the genus Kapala Cameron () (Murray & Heraty 2016) and a similar event would explain the disjunct distribution of Chromoteleia. Alternatively, the dispersal event could have occurred in the opposite direction, followed by radiation in the New World tropics. There is no direct evidence that the distribution of Chromoteleia represents relictual populations but this is likely the case with other scelionine taxa. Archaeoteleia Masner, which today is known from New Zealand and South America, has been documented from (Talamas et al. 2017), suggesting that the extant fauna of this genus is the remainder of a once widespread distribution. Romilius (Chile), Bracalba (Australia) and a species known to us from Baltic amber are considered by some of the authors to be congeneric and the sister taxon to Chromoteleia. If correct, this supports that extinction is the mechanism of a disjunct distribution for a genus equal in age to Chromoteleia.

Character discussion

Basiconic sensilla on A12 Seven of the twenty-seven species of Chromoteleia clearly have two basiconic sensilla on the apical antennomere (Fig. 3.1B), a state that is unknown to us from any other scelionine in which basiconic sensilla are arranged in longitudinal pairs.

Metapleural setation

226 The metapleuron in Chromoteleia is an important source of characters. In all species there is a line of setigerous foveae along the anterior margin of the metapleuron, corresponding to the dorsal portion of the metapleural sulcus. Directly posterior to this line of setae in the dorsal portion of the sclerite (dorsal metapleural area) there may be patch of setae (Fig. 3.4B, black arrow), and its presence or absence separates numerous species. In the ventral portion of the sclerite, there is often a line of setae (Fig. 15, white arrow) directly dorsal to the metapleural triangle. These setae can be distinguished from setation of the metapleural triangle because they do not have foveate bases and are located dorsal to the metapleural epicoxal carina, when this carina is present.

Setation of speculum and subalar pit The presence of setae (Figs 3.4A, 3.4B, blue arrow) on the dorsal speculum and surrounding the subalar pit is found in all species of Chromoteleia and serves as a useful character for generic diagnosis (http://specimage.osu.edu/getImageInfo.html?image_id=89044). However, because this is a newly recognized character, it has yet to be examined thoroughly throughout Scelioninae. This character is known to us from Romilius, Bracalba, and Macroteleia pilosa Muesebeck.

Body color Ashmead presumably coined the name Chromoteleia in reference to the remarkable metallic blue head and mesosoma in the type species. This turns out to be unique within the genus, but most species are colorful in a different way. Only a small number have the body entirely black or dark brown as is generally typical for other scelionines. Most have more or less extensive parts of the mesosoma colored a rusty red to orange. Masner & Hanson (2006) noted that this color pattern is commonly found in a wide range of scelionine genera in species of moderate size (3–10 mm in length) found at lower altitudes (under 2000 m). They attributed the apparent convergent evolution of the color in these taxa to aposematism, a hypothesis that has not yet been critically tested.

World species of Chromoteleia Ashmead C. aequalis Chen & Johnson, sp. n.

227 C. alternata Chen & Johnson, sp. n. C. bidens Chen & Masner, sp. n. C. congoana (Risbec) C. connectens Kieffer = C. brevitarsis Kieffer C. copiosa Chen & Johnson, sp. n. C. cuneus Chen & Johnson, sp. n. C. curta Chen & Johnson, sp. n. C. depilis Chen & Johnson, sp. n. C. dispar Chen & Masner, sp. n. C. feng Chen & Johnson, sp. n. C. fossa Chen & Johnson, sp. n. C. fuscicornis Kieffer C. ingens Chen & Masner, sp. n. C. levitas Chen & Johnson, sp. n. C. longitarsis Kieffer C. longa Chen & Johnson, sp. n. C. maura Chen & Masner, sp. n. C. parvitas Chen & Johnson, sp. n. C. pilus Chen & Johnson, sp. n. C. plana Chen & Johnson, sp. n. C. rara Chen & Johnson, sp. n. C. robusta Chen & Johnson, sp. n. C. rufithorax Kieffer C. semicyanea Ashmead C. semilutea Chen & Johnson, sp. n. C. sparsa Chen & Johnson, sp. n. C. tricarinata Kieffer

Key to Females

228 1 Basiconic sensilla on A7 absent (Figs 3.1B, 3.1F) ...... 2 - Basiconic sensilla on A7 present (Figs 3.1D, 3.2B) ...... 14 2 Median mesoscutal carina absent (Figs 3.35D, 3.40D, 3.41D, 3.42A) ...... 3 - Median mesoscutal carina present (Figs 3.7D, 3.8D, 3.12D, 3.13D, 3.18D, 3.20D, 3.24D, 3.25D, 3.26D, 3.27D, 3.28D, 3.31D, 3.37D, 3.42B, 3.42C) ...... 4 3 Apex of T6 acute in dorsal view (Fig. 3.35F); horn on T1 absent (Fig. 3.35F); median metascutellar carina indistinguishable from surrounding rugae ...... Chromoteleia robusta Chen & Johnson, sp. n. - Apex of T6 rounded in dorsal view (Fig. 3.41F); horn on T1 present (Fig. 3.41F); median metascutellar carina distinct ...... Chromoteleia tricarinata Kieffer 4 A12 with 1 basiconic sensillum ...... 5 - A12 with 2 basiconic sensilla ...... 10 5 Posterior notaulus foveate (Figs 3.12D, 3.13D, 3.26D, 3.27D, 3.28D) ...... 6 - Posterior notaulus smooth (Figs 3.8D, 3.20D, 3.25D) ...... 8 6 Hind basitarsus as long as remaining segments combined; dorsal metapleural area without setae (Figs 3.4A, 3.12B, 3.13B, 3.28B); ventral metapleural area smooth posteriorly (Figs 3.4A, 3.12B, 3.13B, 3.28B) ...... 7 - Hind basitarsus distinctly longer than remaining segments combined (Fig. 3.6F); dorsal metapleural area setose (Fig. 3.4B); ventral metapleural area rugose posteriorly (Fig. 15) ...... Chromoteleia longitarsis Kieffer 7 T1 with horn (Figs 3.12D, 3.13D); occipital carina complete (Figs 3.12D, 3.13D); A6 as long as wide (Fig. 3.13D) ...... Chromoteleia connectens Kieffer - T1 without horn (Fig. 3.28F); occipital carina interrupted medially (Fig. 3.28D); A6 distinctly longer than wide (Fig. 3.1E) ...... Chromoteleia longa Chen & Johnson, sp. n. 8 Apex of T6 rounded in dorsal view (Figs 3.8F, 3.25F); T1 with horn (Figs 3.8F, 3.25F) .....9 - Apex of T6 acute in dorsal view (Fig. 3.20F); T1 without horn (Fig. 3.20F) ...... Chromoteleia feng Chen & Johnson, sp. n. 9 Metasoma variably patterned in alternating orange-yellow and dark brown (Fig. 3.8F); mesoscutellum smooth medially, densely punctate laterally (Fig.3.8D) ...... Chromoteleia alternata Chen & Johnson, sp. n. - Metasoma entirely black (Fig. 3.25F); mesoscutellum densely punctate rugose (Fig. 3.25D)

229 ...... Chromoteleia levitas Chen & Johnson, sp. n. 10 Dorsal metapleural area setose (Figs 3.24B, 3.31B) ...... 11 - Dorsal metapleural area without setae (Figs 3.7B, 3.18B, 3.37B) ...... 12 11 A6 distinctly longer than wide (Fig. 3.24C); occipital carina complete (Fig. 3.24D); hind basitarsus distinctly longer than remaining segments combined ...... Chromoteleia ingens Chen & Masner, sp. n. - A6 as long as wide (Fig. 3.31A); occipital carina interrupted medially (Fig. 3.31D); hind basitarsus as long as remaining segments combined (Fig. 3.31A) ...... Chromoteleia pilus Chen & Johnson, sp. n. 12 Mesosoma black (Fig. 3.7D); netrion densely punctate (Fig. 3.7B); mesoscutal midlobe densely punctate (Fig. 3.7D) ...... Chromoteleia aequalis Chen & Johnson, sp. n. - Mesosoma orange or variably orange to black (Figs 3.18B, 3.18D, 3.37B); netrion longitudinally striate (Figs 3.18B, 3.37B); mesoscutal midlobe densely punctate rugose (Figs 3.18D, 3.37D) ...... 13 13 Occiput rugose (Fig. 3.18D); area directly dorsal to the metapleural triangle with setae (Fig. 3.18B); hind coxa densely punctate (Fig. 3.18B); T6 at least 1.5x longer than wide (Fig. 3.5D) ...... Chromoteleia depilis Chen & Johnson, sp. n. - Occiput smooth (Fig. 3.37D); area directly dorsal to the metapleural triangle without setae (Fig. 3.37B); hind coxa largely smooth, with sparse fine punctures (Fig. 3.37B); T6 approximately as long as wide (Fig. 3.37F) ...... Chromoteleia semilutea Chen & Johnson, sp. n. 14 Apex of T6 acute in dorsal view (Figs 3.5E, 3.15F, 3.29F) ...... 15 - Apex of T6 rounded in dorsal view (Figs 3.6A, 3.11F, 3.14F, 3.16F, 3.17F, 3.19F, 3.21F, 3.30F, 3.32F, 3.33F, 3.36F, 3.38F, 3.39F) ...... 17 15 Mesosoma variably orange to black (Figs 3.9B, 3.15B); netrion longitudinally striate (Figs 3.9B, 3.15B); notaulus foveate (Figs 3.9D, 3.15D) ...... 16 - Mesosoma black (Figs 3.29B, 3.29D); netrion rugose (Fig. 3.29B); notaulus smooth (Fig. 3.29D) ...... Chromoteleia maura Chen & Masner, sp. n. 16 Metascutellum with deeply incised apex, forming two spines laterally (Fig. 3.9D); postmarginal vein approximately as long as stigma vein (Fig. 3.9F); T1 without horn (Fig. 3.9D) ...... Chromoteleia bidens Chen & Masner, sp. n.

230 - Metascutellum trapezoidal with broad apex (Fig. 3.15D); postmarginal vein distinctly shorter than stigma vein (Fig. 3.15F); T1 with horn (Fig. 3.15D) ...... Chromoteleia cuneus Chen & Johnson, sp. n. 17 Hind basitarsus as long as remaining segments combined ...... 18 - Hind basitarsus distinctly longer than remaining segments combined ...... 20 18 Metasoma orange (Fig. 3.19F); mesoscutal midlobe with two rows of foveate grooves along median mesoscutal carina anteriorly, smooth at posterior margin (Fig. 3.19D); postmarginal vein distinctly longer than stigmal vein ...... Chromoteleia dispar Chen & Masner, sp. n. - Metasoma black (Figs 3.30F, 3.38F); mesoscutal midlobe punctate rugose anteriorly, sparsely punctate at posterior margin (Figs 3.30D, 3.38D); postmarginal vein approximately as long as stigmal vein (Figs 3.30F, 3.38F)...... 19 19 A5 as wide as long (Fig. 3.1C); T1 without horn (Fig. 3.30F); T6 approximately as long as wide (Fig. 3.30F) ...... Chromoteleia parvitas Chen & Johnson, sp. n. - A5 distinctly longer than wide (Fig. 3.38B); T1 with horn (Fig. 3.38D); T6 at least 1.5x longer than wide (Fig. 3.38F) ...... Chromoteleia sparsa Chen & Johnson, sp. n. 20 Postmarginal vein distinctly shorter than stigmal vein (Figs 3.10B, 3.22C, 3.23F, 3.33F) .21 - Postmarginal vein as long as or distinctly longer than stigmal vein (Figs 3.14F, 3.16F, 3.21F, 3.32F, 3.36F) ...... 23 21 Frons directly above interantennal process punctate rugose (Fig. 3.11E); dorsal frons with granulate microsculpture (Fig. 3.11E); netrion punctate rugose anteriorly, smooth posteriorly (Fig. 3.11B) ...... Chromoteleia congoana (Risbec) - Frons directly above interantennal process foveolate (Figs 3.22D, 3.23E, 3.33E); dorsal frons without granulate microsculpture (Figs 3.22D, 3.23E, 3.33E); netrion transversely striate (Fig. 3.23B, 3.33B) ...... 22 22 Dorsal metapleural area setose (Fig. 3.23B); mesoscutal midlobe sparsely punctate anteriorly (Fig. 3.23D); T6 sinuate in lateral view ...... Chromoteleia fuscicornis Kieffer - Dorsal metapleural area without setae (Fig. 3.33B); mesoscutal midlobe punctate rugose anteriorly (Fig. 3.33D); T6 flat in lateral view ...... Chromoteleia rara Chen & Johnson, sp. n.

231 23 Head metallic blue (Figs 3.36D, 3.36E); postmarginal vein distinctly longer than stigma vein (Fig. 3.36F); frons without central keel (Fig. 3.36E) ...... Chromoteleia semicyanea Ashmead - Head black (Figs 3.14E, 3.21E, 3.32E); postmarginal vein approximately as long as stigma vein (Figs 3.14F, 3.16F, 3.21F); frons with central keel (Figs 3.14E, 3.16E, 3.21E, 3.32E) ...... 24 24 Mesoscutal midlobe smooth posteriorly (Fig. 3.32D); T2–T3 with a narrow smooth strip medially (Fig. 3.32F) ...... Chromoteleia plana Chen & Johnson, sp. n. - Mesoscutal midlobe sparsely punctate posteriorly (Figs 3.14D, 3.16D, 3.21D); T2–T3 without a narrow smooth strip medially (Figs 3.14F, 3.16F, 3.21F) ...... 25 25 Notaulus foveate (Figs 3.14D, 3.16D); mesepisternum below femoral depression without striae (Figs 3.14B, 3.16B)...... 26 - Notaulus smooth (Fig. 3.21D); mesepisternum below femoral depression with striae (Fig. 3.21B) ...... Chromoteleia fossa Chen & Johnson, sp. n. 26 Netrion transversely striae (Fig. 3.14B); T6 at least 1.5x longer than wide (Fig. 3.14F) ...... Chromoteleia copiosa Chen & Johnson, sp. n. - Netrion rugose (Fig. 3.16B); T6 approximately as long as wide (Fig. 3.16F)...... Chromoteleia curta Chen & Johnson, sp. n.

Key to Males (unknown for C. maura, C. plana, C. fossa, C. parvitas)

1 T7 posterior margin without lateral spines or rounded projections (Fig. 3.6C) ...... 2 - T7 posterior margin with lateral spines (Fig. 3.6E) or rounded projections (Fig. 3.6D) .....14 2 Hind basitarsus as long as remaining segments combined (Fig. 3.6G) ...... 3 - Hind basitarsus distinctly longer than remaining segments combined (Fig. 3.6F) ...... 9 3 Dorsal metapleural area setose (Figs 3.31B, 3.41B) ...... 4 - Dorsal metapleural area without setae (Figs 3.8B, 3.18B, 3.20B, 3.35B, 3.37B) ...... 5 4 Median mesoscutal carina present anteriorly (Fig. 3.31D); ventral metapleural area rugose (Fig. 3.31B) ...... Chromoteleia pilus Chen & Johnson, sp. n. - Median mesoscutal carina absent (Figs 3.40D, 3.41D); ventral metapleural area obliquely striate posteriorly (Fig. 3.41B) ...... Chromoteleia tricarinata Kieffer

232 5 Area directly dorsal to the metapleural triangle with setae (Fig. 3.18B) ...... Chromoteleia depilis Chen & Johnson, sp. n. - Area directly dorsal to the metapleural triangle without setae (Fig. 3.8B, 3.20B, 3.35B, 3.37B) ...... 6 6 Frons with central keel developed only in ventral portion of frons (Figs 3.8E, 3.20E) ...... 7 - Frons with central keel complete, extending dorsally to median ocellus (Figs 3.35E, 3.37E)8 7 Ventral metapleural area obliquely striate posteriorly (Fig. 3.8B); metasoma dark brown to black (Fig. 3.8F) ...... Chromoteleia alternata Chen & Johnson, sp. n. - Ventral metapleural area smooth posteriorly (Fig. 3.20B); metasoma entirely black (Fig. 3.20F)...... Chromoteleia feng Chen & Johnson, sp. n. 8 Metasoma entirely black (Fig. 3.35F); median mesoscutal carina absent (Fig. 3.35D) ...... Chromoteleia robusta Chen & Johnson, sp. n. - Metasoma with T1–T3 orange to dark brown (Fig. 3.37F); median mesoscutal carina present (Fig. 3.37D) ...... Chromoteleia semilutea Chen & Johnson, sp. n. 9 Dorsal A1 striate ...... 10 - Dorsal A1 smooth or punctate ...... 11 10 Mesosoma entirely black (Fig. 3.7D); netrion densely punctate (Fig. 3.7B); dorsal metapleural area without setae (Fig. 3.7B) ...... Chromoteleia aequalis Chen & Johnson, sp. n. - Mesosoma variably orange and black (Fig. 3.27B); netrion rugose (Fig. 3.27B); dorsal metapleural area setose (Fig. 3.4B) ...... Chromoteleia longitarsis Kieffer 11 Head black (Figs 3.14E, 3.16E, 3.33E); frons with central keel (Figs 3.14E, 3.16E, 3.33E); mesopleural carina present (Figs 3.14B, 3.16B, 3.33B) ...... 12 - Head metallic blue (Fig. 3.36E); frons without central keel (Fig. 3.36E) ...... Chromoteleia semicyanea Ashmead 12 Occipital carina interrupted medially (Figs 3.14D, 3.16D); postmarginal vein approximately as long as stigma vein (Figs 3.14F, 3.16F) ...... 13 - Occipital carina complete (Fig. 3.33D); postmarginal vein distinctly shorter than stigma vein (Fig. 3.33F) ...... Chromoteleia rara Chen & Johnson, sp. n. 13 Netrion transversely striate (Fig. 3.14B); epicoxal lobe posterior of propleural epicoxal sulcus smooth ...... Chromoteleia copiosa Chen & Johnson, sp. n.

233 - Netrion rugose (Fig. 3.16B); epicoxal lobe posterior of propleural epicoxal sulcus densely punctate...... Chromoteleia curta Chen & Johnson, sp. n. 14 Dorsal metapleural area setose (Figs 3.9B, 3.23B, 3.24B)...... 15 - Dorsal metapleural area without setae (Figs 3.4A, 3.11B, 3.15B, 3.19B, 3.25B, 3.28B, 3.38B, 3.39B) ...... 17 15 Metascutellum with deeply incised apex, forming two spines laterally (Fig. 3.9D); apex of T7 bispinose (Fig. 3.6E) ...... Chromoteleia bidens Chen & Masner, sp. n. - Metascutellum trapezoidal with broad posterior margin (Figs 3.22B, 3.23D, 3.24D); apex of T7 emarginate between rounded projections (Fig. 3.6D) ...... 16 16 Postmarginal vein distinctly shorter than stigma vein (Fig. 3.23F); A6 as long as wide ...... Chromoteleia fuscicornis Kieffer - Postmarginal vein distinctly longer than stigma vein (Fig. 3.24F); A6 approximately 2.0x longer than wide (Fig. 9) ...... Chromoteleia ingens Chen & Masner, sp. n. 17 Dorsal A1 striate ...... 18 - Dorsal A1 smooth or punctate ...... 20 18 Notaulus foveate (Figs 3.12D, 3.13D, 3.28D); postmarginal vein distinctly longer than stigmal vein (Figs 3.12F, 3.28F) ...... 19 - Notaulus mostly smooth (Fig. 3.25D); postmarginal vein approximately as long as stigmal vein (Fig. 3.25F) ...... Chromoteleia levitas Chen & Johnson, sp. n. 19 Occipital carina complete (Fig. 3.12D); lateral occiput rugose (Fig. 3.12D); gena punctate rugose ventrally (Fig. 3.12B)...... Chromoteleia connectens Kieffer - Occipital carina interrupted medially (Fig. 3.28D); occiput smooth (Fig. 3.28D); gena narrowly smooth ventrally (Fig. 3.28B) ...... Chromoteleia longa Chen & Johnson, sp. n. 20 Frons without central keel (Fig. 3.19E); mesosoma black (Fig. 3.19D); postmarginal vein distinctly longer than stigmal vein (Fig. 3.19F) ...... Chromoteleia dispar Chen & Masner, sp. n. - Frons with central keel (Figs 3.11E, 3.15E, 3.16E); mesosoma variably orange to black (Figs 3.11D, 3.15B, 3.16B); postmarginal vein distinctly shorter than stigmal vein (Figs 3.10B, 3.15F, 3.16F) ...... 21 21 Mesoscutellum sparsely punctate (Fig. 3.11D); T2 without transverse sulcus (Fig. 3.11F); Hind basitarsus distinctly longer than remaining segments combined ......

234 ...... Chromoteleia congoana (Risbec) - Mesoscutellum smooth medially, densely punctate laterally (Figs 3.15D, 3.16D); T2 with transverse sulcus (Figs 3.15F, 3.16F); hind basitarsus as long as remaining segments combined ...... 22 22 Lateral lobe of mesoscutum punctate rugose (Fig. 3.15D); netrion transversely striate (Fig. 3.15B); hind coxa densely punctate (Fig. 3.15B) ...... Chromoteleia cuneus Chen & Johnson, sp. n. - Lateral lobe of mesoscutum sparsely punctate (Fig. 3.16D); netrion rugose (Fig.3.16B); hind coxa largely smooth, with sparse fine punctures ...... Chromoteleia sparsa Chen & Johnson, sp. n.

Species descriptions

Chromoteleia aequalis Chen & Johnson, sp. n. http://zoobank.org/A91D1FE4-DEF0-4D3F-BC48-3F29A67FC42A http://bioguid.osu.edu/xbiod_concepts/452211 Figures 3.7A–3.7F

Description. Body length of female: 6.88–7.27 mm (n=3). Body length of male: 5.90–6.20mm (n=3). Color of A1: yellow to orange. A6 in female: distinctly longer than wide. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 2. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: transversely striate to rugose. Central keel: present, interrupted medially. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: dorsoventrally strigose. Color of mesosoma: black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: densely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: densely punctate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: densely punctate.

235 Macrosculpture of lateral lobe of mesoscutum: densely punctate. Sculpture of notaulus: foveate. Notaular foveae: discrete. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: flat. Sculpture of mesoscutellum: densely punctate rugose. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: areolate. Sculpture of lateral propodeal area: rugose. Mesopleural carina: absent. Sculpture of mesepisternum below femoral depression: smooth directly below femoral depression, otherwise densely punctate. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose anteriorly, smooth posteriorly. Color of legs: pale yellow with tarsi dark brown to black. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: longitudinally punctate rugose. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: smooth to coriaceous. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely punctate to punctate rugose. Diagnosis. This species is easily recognized by its entirely black body both in female and male, densely punctate mesoscutum, and abbreviated median mesoscutal carina. Etymology. The epithet is inspired by the Latin word for equal, in reference to the black body color shared by males and females, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452211] Material Examined. Holotype, female: DOMINICA: Saint Paul Parish, Springfield, 94/017, Archbold Tropical Research Center (ATRC), 27.V.1994, J. B. Woolley, OSUC 221942 (deposited in TAMU). Paratypes: (2 females, 3 males) DOMINICA: 1 female, 3 males, OSUC 584722–584725 (TAMU). GUYANA: 1 female, OSUC 215798 (BPBM).

236 Chromoteleia alternata Chen & Johnson, sp. n. http://zoobank.org/5BA6E119-AF80-47BA-9BF6-A28039A09E23 http://bioguid.osu.edu/xbiod_concepts/318205 Figures 3.8A–3.8F

Description. Body length of female: 5.38–5.90 mm (n=10). Body length of male: 4.40–4.93mm (n=20). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: transversely striate to rugose. Central keel: present only in ventral portion of frons. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: present. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: dorsoventrally strigose. Color of mesosoma: orange. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: sparsely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: rugulose. Sculpture of netrion: punctate rugose anteriorly, smooth posteriorly. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly, sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: smooth. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: rugose anteriorly, sparsely punctate posteriorly. Sculpture of dorsal metapleural area: rugose; smooth. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose anteriorly, obliquely striate posteriorly. Color of legs: orange yellow throughout. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures. Length of postmarginal vein: approximately as long as stigmal vein.

237 Color of metasoma in female: variably patterned in alternating orange yellow and dark brown. Color of metasoma in male: dark brown to black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: densely punctate. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species can be easily distinguished from other Chromoteleia species by the following combination of characters: female metasoma variably patterned in alternating orange yellow and dark brown, central keel present only in ventral portion of frons, and notaulus smooth. Etymology. The epithet is inspired by the Latin word for alternate, in reference to the variably patterned in alternating orange yellow and yellow on metasoma, and is intended to be treated as a noun. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452211] Material Examined. Holotype, female: BRAZIL: ES, Santa Teresa, 660m, 19°58'19"S 40°32'00"W, Santa Lucia Biological Station, 30.V–2.VI.1997, yellow pan trap, W. A. Hoffmann & R. Ribeiro, OSUC 136573 (deposited in OSUC). Paratypes: BRAZIL: 9 females, 56 males, OSUC 149637, 149645, 202506–202508 (AEIC); OSUC 586304–586305, 586307, 586309, 586320–586321, 586323–586326, 586575–586576, 586582–586584, 586588–586589, 586591– 586595, 586597, 586599, 586601–586602 (CNCI); OSUC 199607 (FSCA); OSUC 127206, 128954, 136099, 137616, 148096, 149994, 233157, 233159–233160, 233198, 233332, 235222, 254872–254874, 79997 (MZSP); OSUC 122256, 126972, 138446, 147578, 148058, 148097, 149921, 149995, 150331, 233158, 233161, 233197, 233199, 233331, 246487, 254871, 322558 (OSUC).

Chromoteleia bidens Chen & Masner, sp. n. http://zoobank.org/9FEF89BB-7DA6-461B-BA7B-3B97EE128F9B

238 http://bioguid.osu.edu/xbiod_concepts/318198 Figures 3.2D, 3.5E–3.5F, 3.6E, 3.6H, 3.9A–3.9F

Description. Body length of female: 4.20–4.80 mm (n=16). Body length of male: 4.30–4.67 mm (n=17). Color of A1: yellow to orange. A6 in female: distinctly wider than long. A5 in female: distinctly longer than wide. A6 in male: as long as wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate. Color of head: black. Sculpture of frons directly above interantennal process: transversely striate to rugose. Central keel: present, interrupted medially. Ventral margin of clypeus: straight. Granulate microsculpture of dorsal frons: present. Occipital carina: complete. Granulate microsculpture of vertex: present. Sculpture of occiput: smooth. Sculpture of gena: coarsely punctate rugose. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: sparsely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: sparsely punctate. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: granulate. Shape of metascutellum: elongate trapezoidal but with deeply incised apex, forming two spines laterally. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: absent. Sculpture of mesepisternum below femoral depression: punctate rugose. Sculpture of dorsal metapleural area: rugose; obliquely striate. Setation of dorsal metapleural area: present. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow to brown, with tarsi darker. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: approximately as long as stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: absent. Striae of posterior margin of T1 in female: sparse. Striae of T1 in male: sparse. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate

239 rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: approximately as long as wide. Shape of T6 in female in lateral view: sinuate. Apical spine on female T6: present. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: smooth anteriorly, rugulose posteriorly. Posterior margin of T7 in male: bispinose. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is easily recognized by its bispinose metascutellum a character not found in any other species of Chromoteleia. Etymology. The epithet is inspired by the Latin word for two-toothed, in reference to the bispinose metascutellum, and is intended to be treated as a noun. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=318198] Material Examined. Holotype, female: BRAZIL: RO, 62km S Ariquemes, Rancho Grande Farm, 12.XI–22.XI.1991, E. M. Fisher, OSUC 577455 (deposited in CNCI). Paratypes: (15 females, 15 males) BOLIVIA: 1 female, OSUC 577440 (CNCI). BRAZIL: 8 females, 12 males, OSUC 577452–577453, 577456–577458, 584325 (CNCI); OSUC 204986, 204991 (UCDC); OSUC 185665–185676 (USNM). ECUADOR: 1 female, OSUC 577454 (CNCI). FRENCH GUIANA: 2 females, OSUC 586452, 586827 (CNCI). GUYANA: 2 females, OSUC 232995 (BPBM); OSUC 577451 (CNCI). PERU: 1 female, 3 males, OSUC 577459, 584310, 586806 (CNCI); OSUC 323993 (OSUC).

Chromoteleia congoana (Risbec) http://zoobank.org/381B96B2-0567-4A2C-B2C2-F8E69D0E32A0 http://bioguid.osu.edu/xbiod_concepts/4211 Figures 3.10A–3.10F

Oxyscelio congoana Risbec, 1950: 613 (original description). Chromoteleia congoana (Risbec): Masner, 1976: 25 (description, generic transfer, type information); Johnson, 1992: 364 (cataloged, type information).

Description. Body length of female: 4.38–5.66 mm (n=20). Body length of male: 4.10–4.78 mm (n=4). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic

240 sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate; smooth. Color of head: black. Sculpture of frons directly above interantennal process: punctate rugose. Central keel: complete, extending from interantennal process to median ocellus. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: present. Occipital carina: complete. Granulate microsculpture of vertex: present. Sculpture of occiput: rugose. Sculpture of gena: punctate rugose dorsally and ventrally, strigose medially. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: sparsely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: punctate rugose anteriorly, smooth posteriorly. Microsculpture of mesoscutum: coriaceous. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly, sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: sparsely punctate. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: punctate throughout. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow throughout. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly shorter than stigmal vein. Color of metasoma in female: dark brown. Color of metasoma in male: dark brown to black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: sparse. Transverse sulcus on T2: absent. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: smooth anteriorly, rugulose posteriorly. Posterior margin of T7 in male: emarginate medially between

241 rounded projections. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species can be distinguished from other Chromoteleia species by the following combination of characters: median mesoscutal carina only present anteriorly, mesosoma variably orange to black, postmarginal vein distinctly shorter than stigma vein, T2 without transverse sulcus, male T7 with posterior margin deeply emarginated between rounded projections. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=4211] Material Examined. Holotype, male, Oxyscelio congoana: GABON: Estuaire Prov., Libreville, 1901, C. Chalot, EY11376 (deposited in MNHN). Other material: (35 females, 4 males) CAMEROON: 3 females, 1 male, OSUC 584461–584464 (CNCI). CENTRAL AFRICAN REPUBLIC: 9 females, 1 male, OSUC 226055–226056, 254563, 320836–320837 (OSUC); OSUC 176093, 180933, 207829, 223628, 267420 (SAMC). DEMOCRATIC REPUBLIC OF THE CONGO: 1 male, OSUC 584460 (CNCI). GABON: 1 female, 1 male, OSUC 584457– 584458 (CNCI). GHANA: 1 female, OSUC 412136 (OSUC). GUINEA: 1 female, OSUC 584459 (CNCI). NIGERIA: 20 females, OSUC 584452–584456, 584465–584479 (CNCI). Comments. This species is only found in Afrotropical region.

Chromoteleia connectens Kieffer http://zoobank.org/7409F8CA-0594-4232-8481-FCC20CDC290E http://bioguid.osu.edu/xbiod_concepts/4212 Figures 3.4A, 3.12A–3.12F, 3.42B

Chromoteleia (Oxyscelio) connectens Kieffer, 1910a: 313 (original description, keyed). Oxyscelio (Oxyscelio) connectens (Kieffer): Kieffer, 1910b: 69 (generic transfer). Oxyscelio connectens (Kieffer): Kieffer, 1926: 361, 362 (description, keyed); Dodd, 1931: 77 (excluded from Oxyscelio, position uncertain). Chromoteleia connectens Kieffer: Masner, 1976: 25 (type information); Hoebeke, 1980: 26 (type information); Johnson, 1992: 364 (cataloged, type information).

242 Chromoteleia brevitarsis Kieffer, 1910a: 313, 315 (original description, keyed); Masner, 1976: 25 (type information, junior synonym of Chromoteleia connectens Kieffer); Johnson, 1992: 364 (type information). Petalosema brevitarsis (Kieffer): Kieffer, 1926: 358, 360 (generic transfer, description, keyed). http://zoobank.org/C7398CF1-267D-4D5D-B89C-55697354254C http://bioguid.osu.edu/xbiod_concepts/8526

Description. Body length of female: 5.38–5.95 mm (n=20). Body length of male: 5.10–5.86 mm (n=20). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: transversely striate to rugose. Central keel: complete, extending from interantennal process to median ocellus. Ventral margin of clypeus: straight. Granulate microsculpture of dorsal frons: present. Occipital carina: complete. Granulate microsculpture of vertex: absent; present. Sculpture of occiput: rugose. Sculpture of gena: dorsoventrally strigose. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: rugose. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose throughout. Macrosculpture of lateral lobe of mesoscutum: punctate rugose. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present along full length of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely punctate laterally; longitudinally carinate medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose; smooth. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: rugose anteriorly, sparsely punctate posteriorly. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose anteriorly, smooth posteriorly. Color of

243 legs: orange yellow throughout; orange to pale brown. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: black; mostly black with T1–T3 orange to dark brown. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: sparse. Striae of T1 in male: sparse. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely punctate and granulate. Length of T6 in female: approximately as long as wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: granulate. Posterior margin of T7 in male: emarginate medially between rounded projections. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species can be distinguished from other Chromoteleia by the following combination of characters: female A7 without basiconic sensillum, occipital carina complete, median mesoscutal carina present along full length of mesoscutum, ventral metapleural area smooth posteriorly, postmarginal vein distinctly longer than stigma vein, male T7 with posterior margin deeply emarginated between rounded projections. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=4212] Material Examined. Holotype, female, C. brevitarsis: BRAZIL: PA, no date, Baker, CAS TYPE9597 (deposited in CAS). Holotype, female, C. connectens: BRAZIL: PA, no date, Baker, CAS TYPE9618 (deposited in CAS). Other material: (524 females, 594 males) BELIZE: 1 female, OSUC 586444 (CNCI). BOLIVIA: 5 females, 7 males, OSUC 584139–584140, 584298–584300 (CNCI); DPI_FSCA 00010202–00010208 (CSCA). BRAZIL: 77 females, 73 males, OSUC 202485, 202488, 202490, 202493, 202495–202497, 202499–202501, 202515– 202517, 202519, 202523, 202532, 202535 (AEIC); OSUC 232991 (BPBM); OSUC 583467– 583468, 583471, 583473–583475, 583484–583485, 583492, 583495–583496, 583498, 583510, 583516, 584146, 584152, 584161, 584164–584165, 584171–584175, 584177, 584330, 584334, 584336, 586124–586126, 586131–586132, 586134, 586352, 586356, 586358, 586365, 586368, 586394–586395, 586397–586398, 586404, 586406, 586418–586419, 586567–586568, 586796 (CNCI); OSUC 199611, 199634 (FSCA); OSUC 181648, 225221, 225223, 225240, 231875, 233121, 233201, 233295, 233345, 235221, 267216, 322173, 322599, 326391, 326465, 344484,

244 354669, 359051, 359057, 359059, 359065, 359067, 359076, 370978, 379216, 380242, 449172, 449186–449187 (MPEG); OSUC 112474, 112594, 112933, 132283, 132337–132338 (MZSP); OSUC 132223, 132546, 132576, 132614, 225220, 225239, 233200, 233320, 255000, 322166, 322175, 322598, 322639, 326569, 345343, 359056, 359063–359064, 359068, 370976–370977, 379214, 381994, 449168, 449178, 48530, 48535, 56248, 56250, 56257–56258, 56260, 56262 (OSUC); OSUC 204987, 204989 (UCDC); OSUC 225298–225300, 225302–225304, 225309, 225312, 225314–225315 (USNM). COLOMBIA: 49 females, 55 males, OSUC 584137, 584301 (CNCI); OSUC 143884, 143894–143895, 143897, 143899, 143901, 144022, 152042–152043, 152069, 152071, 162521, 162680–162681, 166453, 166456–166457, 166521, 166555, 170459, 176951, 182582–182583, 182713, 188629–188630, 188632–188633, 188691–188692, 188783, 189043–189044, 189046, 190347–190348, 190350, 190901, 191362, 193136, 193185, 193429, 193456–193458, 193460, 193661, 193664, 193680, 193819, 193960–193962, 228571, 228601, 228603–228605, 228607, 230379, 230434, 230436, 230439, 232724, 232726, 249925–249926, 253478, 269375, 270003, 270005–270007, 321676 (IAVH); OSUC 143896, 143898, 152070, 152072, 162520, 162679, 166450, 166458, 166520, 166565, 166588, 170460, 176950, 182581, 189045, 190349, 191043, 191173, 193168, 193184, 193793, 228602, 228606, 232723, 249928, 269377, 270004, 273444 (OSUC). COSTA RICA: 145 females, 195 males, 1 unknown, OSUC 202537, 202539–202540, 202549, 205189 (AEIC); OSUC 232993 (BPBM); OSUC 232189, 253965, 253967–253968, 253970–253971, 556950, 556971, 556973–556977, 556980, 556982– 556985, 556993, 556995–556996, 556999, 557001, 557004, 557010, 557014, 557018, 557023, 557048, 557050–557051, 557053, 557058, 557084, 557087, 577888–577893, 577895–577896, 577899, 577908, 577927, 577929, 577938, 577940, 577943–577944, 577947, 577949–577951, 577954–577955, 577962, 577964, 577971, 577974–577978, 577980–577984, 577986–577987, 577989–577990, 577992–577999, 578001, 578007, 578009, 578012–578013, 578018, 578020– 578022, 578025–578043, 578045, 578047–578052, 578066, 578073, 578076, 578080, 578085, 578092, 578097, 578099, 578104, 583413, 583415–583416, 583422–583423, 583461, 583479, 583482, 583508, 583511–583513, 583731, 583739, 583756, 583760, 583763–583764, 583768– 583771, 583773, 583787, 583794, 583796, 583834, 583850, 583878, 583881, 583890–583892, 583896–583897, 583901, 583910, 583916, 583926–583928, 583931–583938, 583947–583948, 583978, 583980, 583982, 583988–583990, 583992, 583994, 583997–583998, 584003–584006, 584008–584009, 584012, 584014, 584016–584017, 584019, 584023, 584026–584028, 584031,

245 584226, 584228, 584231–584232, 584234, 584237, 584239, 584241, 584245–584246, 584248– 584249, 584251, 584253–584259, 584271–584273, 584277–584280, 584284–584286, 584288– 584293, 586121, 586123, 586130, 586150–586151, 586161, 586163–586164, 586166–586168, 586171, 586179–586181, 586183, 586194, 586196, 586198, 586200–586201, 586203, 586205– 586206, 586209, 586212–586214, 586216, 586260–586261, 586268–586269, 586271–586274, 586293–586295, 586302, 586357, 586390, 586399, 586415, 586449, 586451, 586471, 586474– 586476, 586485, 586493, 586499, 586501, 586510, 586513, 586516–586520, 586541–586543, 586559, 586619–586620, 586638–586640, 586645, 586648–586649, 586656, 586658–586659, 586679, 586681, 586764, 586778, 586781, 586783, 586785–586787, 586831, 586833, 586837, 586839, 586841 (CNCI); DPI_FSCA 00010229–00010230, SM0810076, SM0810559 (KUNH); OSUC 205812 (UCDC). ECUADOR: 122 females, 166 males, OSUC 149644, 202502 (AEIC); OSUC 181428, 557030, 557034, 557037, 557042–557045, 557092–557094, 557096, 557098, 583476–583477, 583751, 584047–584059, 584061–584073, 584075–584082, 584084–584096, 584099–584101, 584105, 584110–584111, 584120–584121, 584123–584130, 584132, 584134– 584136, 584399, 584401, 584403, 584405–584406, 584408, 584411–584413, 584415, 584427, 584430, 584432–584451, 584807–584816, 584818–584837, 584839–584856, 585089–585090, 585097, 586142–586144, 586178, 586311–586313, 586317–586319, 586335, 586359, 586413, 586546–586553, 586555–586556, 586558, 586625, 586633, 586668–586670, 586672–586676, 586693, 586696–586707, 586709–586713, 586715–586720, 586723, 586725–586726, 586728– 586730, 586732–586746, 586873 (CNCI); DPI_FSCA 00010184–00010201, 00010209 (CSCA); OSUC 199615–199629 (FSCA); OSUC 221931, 221939 (TAMU); OSUC 204993, 204995–204996, 204998 (UCDC). FRENCH GUIANA: 36 females, 28 males, OSUC 555804, 555814, 555819, 586222, 586225, 586228–586230, 586232, 586234–586238, 586240–586242, 586244–586245, 586247–586255, 586257, 586423, 586427, 586434, 586436, 586439, 586443, 586446, 586453–586458, 586461–586464, 586609–586612, 586830, 586844–586845, 586850– 586851, 586854–586855, 586859 (CNCI); SM0096678 (KUNH); OSUC 267215, 47019–47021, 47023 (OSUC). GUATEMALA: 2 females, 1 male, OSUC 203118 (AEIC); OSUC 584925 (CNCI); OSUC 204701 (UCDC). HONDURAS: 1 male, OSUC 413764 (MZLU). PANAMA: 8 females, 8 males, OSUC 556951, 584182, 584379, 586623v586624, 586626, 586629, 586824 (CNCI); DPI_FSCA 00010227 (KUNH); OSUC 221921, 233027, 271017–271020, 320637 (TAMU). PERU: 24 females, 17 males, OSUC 149639 (AEIC); OSUC 578057, 578060,

246 578062, 584305, 584307–584308, 584314–584316, 586135–586137, 586185–586188, 586190– 586192, 586421–586422, 586441–586442, 586805, 586808–586814, 586846, 586865 (CNCI); OSUC 218803–218805 (INHS); DPI_FSCA 00010228 (KUNH); OSUC 223890, 323994 (OSUC); OSUC 225399 (USNM). VENEZUELA: 55 females, 43 males, OSUC 149646 (AEIC); OSUC 578015, 583517, 586145 (CNCI); OSUC 230357–230359, 230361, 232281, 232827–232830, 251645–251647, 251649, 251651–251652, 251655, 251657–251661, 251663– 251666, 251668–251670, 251672–251675, 251677–251681, 251683–251684, 251686–251688, 251690, 320743–320745, 320747–320753, 321367, 321369–321371, 321373–321374, 321376– 321379, 321382, 323405, 323407–323412, 323414, 381986–381987, 381989, 381991–381992 (MIZA); OSUC 230360, 232282, 251643, 251648, 251650, 251653, 251656, 251671, 251682, 251685, 251692, 321368, 321372, 321375, 321380–321381, 323406 (OSUC). Comments. This species exhibits variation in color and microsculpture. The development of microsculpture on vertex could be absent or present; mesoscutum smooth or carinate medially; mesoscutellum, mesopleuron, and metapleuron ranging from orange to dark brown or black.

Chromoteleia copiosa Chen & Johnson, sp. n. http://zoobank.org/E61DC4D8-138F-4525-8A8A-79476002AAA8 http://bioguid.osu.edu/xbiod_concepts/452220 Figures 3.14A–3.14F

Description. Body length of female: 4.78–5.76 mm (n=20). Body length of male: 4.30–5.33 mm (n=20). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: as long as wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate. Color of head: black. Sculpture of frons directly above interantennal process: areolate. Central keel: complete, extending from interantennal process to median ocellus. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: punctate rugose dorsally and ventrally, strigose medially. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical

247 sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly, sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: densely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: punctate rugose. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow throughout. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: approximately as long as stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: sparse; dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: coriaceous anteriorly, densely punctate posteriorly. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. sparsa, but it can be distinguished by its medially interrupted occipital carina, transversely striate netrion, and the hind basitarsus that is distinctly longer than the remaining segments combined. Etymology. The epithet is inspired by the Latin word for abundant, in reference to the densely punctate rugose ventral mesepisternum, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452220]

248 Material Examined. Holotype, female: BELIZE: Orange Walk Dist., Rio Bravo Conservation and Management Area, 15.VII–21.VII.1996, Malaise trap, P. Kovarik, OSUC 577518 (deposited in CNCI). Paratypes: (54 females, 10 males) BELIZE: 11 females, OSUC 556953–556954, 556966, 577508, 577511–577512, 577516–577517, 586761 (CNCI); OSUC 254866–254867 (OSUC). BRAZIL: 1 male, OSUC 584162 (CNCI). COLOMBIA: 1 female, OSUC 584304 (CNCI). COSTA RICA: 33 females, 6 males, OSUC 577902, 577910–577911, 577914–577916, 577918–577922, 583913, 583954, 583956, 583958–583959, 583962, 583964–583965, 583967– 583970, 583972–583975, 584033 (CNCI); OSUC 577909, 577913, 577917, 583955, 583957, 583960–583961, 583963, 583966, 583971, 586162 (OSUC). GUATEMALA: 2 females, OSUC 577513, 577520 (CNCI). HONDURAS: 1 male, OSUC 577515 (CNCI). MEXICO: 4 females, 2 males, OSUC 577507, 577514, 577519, 584721 (CNCI); OSUC 271012 (OSUC); OSUC 271015 (UNAM). PANAMA: 1 female, OSUC 584382 (CNCI). TRINIDAD AND TOBAGO: 1 female, OSUC 583478 (CNCI). VENEZUELA: 1 female, OSUC 586289 (CNCI).

Chromoteleia cuneus Chen & Johnson, sp. n. http://zoobank.org/BD48157A-CEA8-45EA-BCC2-11E2807CA094 http://bioguid.osu.edu/xbiod_concepts/452221 Figures 3.3C, 3.15A–3.15F

Description. Body length of female: 4.88–5.10 mm (n=20). Body length of male: 4.30–4.89 mm (n=20). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate; smooth. Color of head: black. Sculpture of frons directly above interantennal process: areolate. Central keel: present only in ventral portion of frons. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: coarsely punctate rugose. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: transversely striate.

249 Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly, sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: punctate rugose. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: largely smooth, punctate rugose anteriorly and directly below femoral depression. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow to brown, with tarsi darker. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly shorter than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: sinuate. Apical spine on female T6: present. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: coriaceous anteriorly, densely punctate posteriorly. Posterior margin of T7 in male: emarginate medially between rounded projections. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species can be distinguished from other Chromoteleia by the following combination of characters: frons with central keel developed only in ventral portion of frons, dorsal metapleural area without setae, postmarginal vein distinctly shorter than stigmal vein, apex of T6 acute in dorsal view, male T7 with posterior margin deeply emarginate medially between rounded projections. Etymology. The epithet is inspired by the Latin word for wedge, in reference to the shape of T6 in female, and is intended to be treated as a noun.

250 Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452221] Material Examined. Holotype, female: ECUADOR: Santo Domingo de los Tsáchilas Prov., 16km SE Santo Domingo de los Colorados, Tinalandia, 500m, 4.VI–14.VI.1976, S. Peck & J. Peck, OSUC 585001 (deposited in CNCI). Paratypes: (61 females, 52 males) COSTA RICA: 2 females, OSUC 577898, 586775 (CNCI). ECUADOR: 58 females, 52 males, OSUC 202533 (AEIC); OSUC 583757, 584074, 584107–584109, 584416–584419, 584421–584422, 584424, 584426, 584758, 585003–585004, 585006–585012, 585014–585021, 585023–585028, 585030– 585043, 585045–585076, 585078, 585080, 585082–585088, 586327–586328, 586635, 586847– 586848 (CNCI); OSUC 369619 (MZLU); OSUC 584420, 584423, 584425, 585002, 585005, 585013, 585022, 585029, 585077, 585079, 585081 (OSUC). PANAMA: 1 female, OSUC 271021 (TAMU). Other material: ECUADOR: 2 females, 1 male, DPI_FSCA 00010213– 00010215 (CSCA).

Chromoteleia curta Chen & Johnson, sp. n. http://zoobank.org/E661A7C8-E5DA-4F9A-8507-6D8447DC59AA http://bioguid.osu.edu/xbiod_concepts/318230 Figures 3.16A–3.16F, 3.42D

Description. Body length of female: 4.23–5.27 mm (n=20). Body length of male: 4.10–5.13 mm (n=20). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: as long as wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate. Color of head: black. Sculpture of frons directly above interantennal process: punctate rugose. Central keel: present, interrupted medially. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: narrowly smooth ventrally, punctate rugose dorsally. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: densely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: rugose. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly,

251 sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: flat. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: punctate throughout. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow throughout. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures. Length of postmarginal vein: approximately as long as stigmal vein. Color of metasoma in female: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: sparse. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: approximately as long as wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: granulate. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. parvitas, but it can be distinguished by the combination of the following characters: A5 in female distinctly longer than wide, postmarginal vein approximately as long as stigmal vein, female T1 horn present. Etymology. The epithet is inspired by the Latin word for shortened, in reference to the short median mesoscutal carina, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=318230] Material Examined. Holotype, female: BRAZIL: SP, Trilha da Anta, Base Barra Grande, YPT B0, Intervales State Park, 11.XII–14.XII.2000, yellow pan trap, M. T. Tavares, OSUC 185626 (deposited in MZSP). Paratypes: (22 females, 25 males) BRAZIL: 13 females, 19 males, OSUC 202509 (AEIC); OSUC 578000, 583491, 586148, 586396 (CNCI); OSUC 126962, 129282,

252 147965, 148090, 148098, 148100, 149925, 149996, 150330, 150442, 185627, 263040, 66311, 79986 (MZSP); OSUC 127095, 128181, 128644, 147683–147684, 147730, 148057, 148059, 148089, 149844, 149997, 185628, 79987 (OSUC). MEXICO: 1 male, OSUC 586432 (CNCI). VENEZUELA: 9 females, 5 males, OSUC 578014, 578016–578017, 584142 (CNCI); OSUC 251654, 251689, 376068, 376070–376071 (MIZA); OSUC 376069, 376072–376073, 376075, 381988 (OSUC). Comments. This species is extremely variable in mesosoma color and can be roughly divided into dark and light forms.

Chromoteleia depilis Chen & Johnson, sp. n. http://zoobank.org/30F7A6B0-982F-4A20-8ACC-EDA6322C9E22 http://bioguid.osu.edu/xbiod_concepts/452216 Figures 3.5C–3.5D, 3.6G, 3.18A–3.18F

Description. Body length of female: 5.48–7.65 mm (n=20). Body length of male: 4.34–5.85mm (n=20). Color of A1: yellow to orange. A6 in female: distinctly longer than wide. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 2. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: transversely striate to rugose. Central keel: complete, extending from interantennal process to median ocellus. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: present. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: rugose. Sculpture of gena: dorsoventrally strigose. Color of mesosoma: variably orange to black; orange. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose throughout. Macrosculpture of lateral lobe of mesoscutum: punctate rugose. Sculpture of notaulus: foveate. Notaular foveae: discrete. Median mesoscutal carina: present along full length of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely

253 punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: absent; present. Sculpture of mesepisternum below femoral depression: smooth directly below femoral depression, otherwise densely punctate. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow to brown, with tarsi darker. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: smooth to coriaceous. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. pilus, but it can be distinguished by the combination of the following characters: dorsal metapleural area without setation, A6 in female distinctly longer than wide. Etymology. The epithet is inspired by the Latin word for hairless, in reference to the hairless dorsal metapleural area, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452216] Material Examined. Holotype, female: GUATEMALA: Zacapa Dept., 3.5km SE La Unión, 1500m, 20.VI–27.VI.1993, flight intercept trap, J. Ashe & R. Brooks, OSUC 577436 (deposited in CNCI). Paratypes: (75 females, 38 males) BELIZE: 11 females, 1 male, OSUC 556955, 583463–583464, 584746–584747, 586420, 586426, 586748–586750, 586759–586760 (CNCI). COSTA RICA: 23 females, 12 males, OSUC 232059, 232187, 253969, 556990–556991, 557062, 577484–577487, 583436, 583889, 583952–583953, 583981, 584042, 584045, 584767–

254 584777, 586487, 586489, 586494, 586500, 586534, 586538 (CNCI); OSUC 369615 (MZLU). EL SALVADOR: 2 females, 1 male, OSUC 583419, 586840, 586842 (CNCI). GUATEMALA: 3 females, 3 males, OSUC 234526 (BPBM); OSUC 577437–577439, 584728–584729 (CNCI). HONDURAS: 5 females, OSUC 199605 (FSCA); OSUC 369623, 369625, 413765, 413772 (MZLU). MEXICO: 31 females, 16 males, OSUC 149648–149649, 202503 (AEIC); OSUC 577496–577498, 584726–584727, 584730–584745, 586428–586431 (CNCI); OSUC 271006– 271010, 321342, 322559, 372145, 377930, 415086–415089, 49325, 576180–576181, 576183 (OSUC); OSUC 221930, 221934 (TAMU). PANAMA: 3 males, OSUC 202560 (AEIC); OSUC 232813, 319210 (TAMU). VENEZUELA: 2 males, OSUC 376076–376077 (MIZA).

Chromoteleia dispar Chen & Masner, sp. n. http://zoobank.org/C811B539-CFDC-4A2E-8EB7-D9664F889DC7 http://bioguid.osu.edu/xbiod_concepts/318239 Figures 3.19A–3.19F

Description. Body length of female: 4.88–5.50 mm (n=20). Body length of male: 4.60–5.29 mm (n=20). Color of A1: yellow to orange. A6 in female: distinctly wider than long. A5 in female: as long as wide. A6 in male: as long as wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate. Color of head: black. Sculpture of frons directly above interantennal process: areolate. Central keel: absent. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth medially, striate laterally. Sculpture of gena: coarsely punctate rugose. Color of mesosoma: black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: densely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: two rows of foveate grooves along median mesoscutal carina anteriorly, smooth at posterior margin. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: densely punctate rugose.

255 Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: areolate. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: punctate rugose. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow throughout. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: orange. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: sparse. Striae of T1 in male: sparse. Transverse sulcus on T2: present. Sculpture of T2: longitudinally punctate rugose. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: approximately as long as wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: smooth to coriaceous. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species can be easily distinguished by the orange female metasoma, the lack of central keel, and the two rows of foveate grooves along median mesoscutal carina anteriorly. Etymology. The epithet is inspired by the Latin word for different, in reference to the different colors of metasoma between female and male, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=318239] Material Examined. Holotype, female: COLOMBIA: Magdalena Dept., Zaino, M.567, 50m, 11°20'N 74°02'W, Tayrona Natural National Park, 28.VII–14.VIII.2000, Malaise trap, R. Henriquez, OSUC 190774 (deposited in IAVH). Paratypes: (22 females, 28 males) COLOMBIA: 11 females, 17 males, OSUC 586769–586770 (CNCI); OSUC 151957–151959, 151961, 152023, 170413, 190080, 190082, 190773 (IAVH); OSUC 151960, 152020, 152022, 170414, 190081, 223860, 223866 (OSUC); OSUC 223857–223859, 223861–223865, 223867– 223868 (USNM). VENEZUELA: 11 females, 11 males, OSUC 577442–577443, 577446– 577448 (CNCI); OSUC 237132, 377646 (MIZA); OSUC 237133–237135, 271022–271028, 577441, 577445, 586469 (OSUC); OSUC 221940 (TAMU); OSUC 577444 (USNM).

256 Chromoteleia feng Chen & Johnson, sp. n. http://zoobank.org/A252B7DB-459E-44BB-97A8-72AC6EC76161 http://bioguid.osu.edu/xbiod_concepts/452223 Figures 3.20A–3.20F

Description. Body length of female: 4.48–5.15 mm (n=20). Body length of male: 4.30–5.12 mm (n=20). Color of A1: yellow to orange. A6 in female: distinctly wider than long. A5 in female: distinctly longer than wide. A6 in male: as long as wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: transversely striate to rugose. Central keel: present only in ventral portion of frons. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: present. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: narrowly smooth ventrally, punctate rugose dorsally. Color of mesosoma: orange. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly, sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: smooth. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: densely punctate rugose. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: smooth. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: striate to densely punctate below mesopleural carina, otherwise smooth. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose anteriorly, smooth posteriorly. Color of legs: orange yellow throughout. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures. Length of postmarginal vein: approximately as long as stigmal vein.

257 Color of metasoma in female: black. Horn on T1 in female: absent. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: absent; present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: approximately as long as wide. Shape of T6 in female in lateral view: sinuate. Apical spine on female T6: present. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: densely punctate. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. robusta in the acute T6 apex, but it can be distinguished by the combination of the following characters: female A12 has only one basiconic sensillum, occipital carina complete, notaulus smooth.

Etymology. The epithet is inspired by the Chinese word 锋 (fēng) for the name of the sharp point of a spear, in reference to the pointed female T6, and is intended to be treated as a noun. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=318239] Material Examined. Holotype, female: BRAZIL: PA, Melgaço Mpio., Ferreira Penna Scientific Station, trail, P05057, 01°44'28"S 51°27'21.3"W, Caxiuanã, 16.XI–19.XI.2003, yellow pan trap, A. P. Aguiar & J. Dias, OSUC 233344 (deposited in MPEG). Paratypes: (24 females, 92 males) BRAZIL: 16 females, 90 males, OSUC 202483, 202487, 202518, 202520–202521, 202526–202527 (AEIC); OSUC 584166–584170, 584176, 586360, 586363–586364, 586366– 586367, 586370, 586372 (CNCI); OSUC 185794, 185832, 185834–185835, 185837, 185839, 185841, 225228–225229, 225233, 231866, 231868, 231870, 231872–231873, 231886, 241252, 241254, 241256, 241279, 322168, 322170, 322172, 322176, 322594, 322596, 322600, 326258, 326260, 359052, 359058, 359066, 359078, 380236–380237, 380240, 449170, 449174, 449177 (MPEG); OSUC 166091, 185792, 185795, 185833, 185836, 185838, 185840, 225222, 225226, 225231, 225238, 231867, 231869, 231871, 231874, 233120, 233293, 241251, 241255, 241257, 254999, 322167, 322169, 322174, 322202, 322595, 322597, 322640, 326259, 359054–359055, 359060, 359070, 359074, 366730, 371944, 380235, 380238, 380241, 381993, 449166, 449173, 449175–449176, 449182 (OSUC); OSUC 225301, 225310 (USNM). FRENCH GUIANA: 7 females, 2 males, OSUC 555821, 586223–586224, 586226–586227, 586231, 586829, 586852, 586858 (CNCI). VENEZUELA: 1 female, OSUC 557019 (CNCI).

258 Chromoteleia fossa Chen & Johnson, sp. n. http://zoobank.org/06CBBAC3-7C54-4A69-9395-ED4B55E67D7D http://bioguid.osu.edu/xbiod_concepts/452226 Figures 3.21A–3.21F

Description. Body length of female: 6.90 mm (n=1). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate; smooth. Color of head: black. Sculpture of frons directly above interantennal process: punctate rugose. Central keel: present only in ventral portion of frons. Ventral margin of clypeus: straight. Granulate microsculpture of dorsal frons: absent. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: dorsoventrally strigose; narrowly smooth ventrally, punctate rugose dorsally. Color of mesosoma: orange. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: densely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly, sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: punctate rugose. Sculpture of notaulus: smooth. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: striate to densely punctate below mesopleural carina, otherwise smooth. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow throughout. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures. Length of postmarginal vein: approximately as long as stigmal vein.

259 Color of metasoma in female: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. feng with which it shares the characters of smooth notaulus and female A12 with one basiconic sensillum, but it can be distinguished by the combination of the following characters: female T6 without apical spine, occipital carina interrupted medially. Etymology. The epithet is inspired by the Latin word for ditch, in reference to the smooth notaulus, and is intended to be treated as a noun. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452226] Material Examined.Holotype, female: PANAMA: Kuna Yala Comarca, 2001/038, 390m, 09°20'31"N 78°59'38"W, Nusagandi Station, 12.I–15.I.2001, yellow pan trap, M. Yoder, OSUC 320582 (deposited in TAMU).

Chromoteleia fuscicornis Kieffer http://zoobank.org/FFE02947-921E-4224-971E-7C0259A61634 http://bioguid.osu.edu/xbiod_concepts/4213 Figures 3.6A–3.6B, 3.6D, 3.22A–3.23F

Chromoteleia fuscicornis Kieffer, 1910a: 313, 316 (original description, keyed); Masner, 1976: 25 (description, type information); Johnson, 1992: 364 (cataloged, type information). Petalosema fuscicornis (Kieffer): Kieffer, 1926: 358, 360 (generic transfer, description, keyed).

Description. Body length of female: 4.28–6.90 mm (n=20). Body length of male: 4.10–6.53 mm (n=20). Color of A1: yellow to orange. A6 in female: distinctly wider than long. A5 in female: distinctly longer than wide; as long as wide. A6 in male: as long as wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate; smooth. Color of head: black. Sculpture of frons directly above interantennal process: areolate.

260 Central keel: complete, extending from interantennal process to median ocellus. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth medially, striate laterally. Sculpture of gena: narrowly smooth ventrally, punctate rugose dorsally. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: sparsely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: sparsely punctate. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: smooth. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: punctate rugose. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: present. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange to pale brown. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly shorter than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: sparse. Transverse sulcus on T2: absent. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: sinuate. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: smooth anteriorly, rugulose posteriorly. Posterior margin of T7 in male: emarginate medially between

261 rounded projections. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species can be distinguished from other Chromoteleia by the following combination of characters: female A7 with one basiconic sensillum, dorsal metapleural area with setae, basitarsus distinctly longer than remaining segments combined, postmarginal vein distinctly shorter than stigmal vein, male T7 with posterior margin deeply emarginate medially between rounded projections. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=4213] Material Examined. Holotype, female: BRAZIL: PA, no date, Baker, CAS TYPE9649 (deposited in CAS). Other material: (164 females, 155 males) BELIZE: 1 female, OSUC 577461 (CNCI). BOLIVIA: 2 females, OSUC 586776–586777 (CNCI). BRAZIL: 45 females, 78 males, OSUC 149647, 202479–202481, 202484, 202486, 202489, 202491, 202494 (AEIC); OSUC 577932, 583469, 583472, 583494, 583497, 583503, 584147–584148, 584150–584151, 584154–584156, 584160, 584324, 584327–584329, 584335, 586129, 586133, 586147, 586303, 586314–586315, 586329, 586369, 586371, 586393, 586400–586403, 586416–586417, 586573, 586788–586795 (CNCI); OSUC 199636 (FSCA); OSUC 185793, 185831, 225230, 225587, 251722, 322601, 326463, 326475, 344379, 354649, 354653, 359053, 359071, 359075, 359077, 379217, 380234, 449167, 449171, 449185 (MPEG); OSUC 185796, 185842, 225224, 225234, 225585, 241249, 322171, 326261, 326464, 326476–326477, 344338, 344377, 354577, 354671, 354679, 359061–359062, 359069, 359072–359073, 366731, 374856, 380239, 380244, 449169, 449180, 449183, 56224, 56226, 56244–56247, 56249, 56251–56256, 56259 (OSUC); OSUC 204890, 204906–204907, 204988, 204992 (UCDC); OSUC 225316–225317 (USNM). COLOMBIA: 23 females, 11 males, OSUC 586306, 586346, 586762 (CNCI); OSUC 143900, 151956, 166454–166455, 166524, 189042, 189623, 190775, 191044, 230435, 232722, 273445, 320929 (IAVH); OSUC 143893, 151955, 166452, 166519, 166523, 166541, 170415, 182584, 190079, 190816, 191293, 193459, 232725, 249927, 267217, 269374, 269376, 321675 (OSUC). COSTA RICA: 3 females, OSUC 556987, 557081, 586411 (CNCI). ECUADOR: 61 females, 38 males, OSUC 149650 (AEIC); OSUC 556943, 557031, 557033, 557035, 557038–557039, 557090, 557095, 583706, 583708, 583753, 584060, 584098, 584102–584103, 584112, 584114– 584117, 584119, 584131, 584133, 584398, 584400, 584404, 584407, 584409, 584414, 584428– 584429, 584431, 584759–584762, 585099–585105, 585107–585108, 585110–585118, 585120,

262 586159–586160, 586177, 586316, 586330, 586334, 586336, 586342, 586412, 586502–586503, 586505, 586554, 586667, 586694–586695, 586708, 586714, 586724, 586731 (CNCI); DPI_FSCA 00010220 (CSCA); OSUC 199630 (FSCA); OSUC 557032, 557041, 557089, 557097, 584083, 584097, 584104, 584113, 584118, 584402, 584410, 585109, 585119, 586331, 586333, 586531, 586727 (OSUC); OSUC 221926–221927 (TAMU); OSUC 204994, 204997 (UCDC). FRENCH GUIANA: 6 females, 4 males, OSUC 586233, 586243, 586246, 586256, 586433, 586828, 586849, 586857, 586862, 586864 (CNCI). PERU: 18 females, 24 males, OSUC 556970, 578055, 578061, 584309, 584311–584313, 584318–584319, 584321–584322, 586138, 586189, 586440, 586572, 586797–586801, 586803–586804, 586866 (CNCI); DPI_FSCA 00010216–00010219, 00010221–00010224 (CSCA); OSUC 199609 (FSCA); DPI_FSCA 00010225–00010226, SM0267120 (KUNH); OSUC 181564, 323995, 584320, 586802, 586807 (OSUC); OSUC 204984–204985 (UCDC). SURINAME: 2 females, OSUC 557085, 584297 (CNCI). VENEZUELA: 3 females, OSUC 586157, 586470, 586472 (CNCI).

Chromoteleia ingens Chen & Masner, sp. n. http://zoobank.org/0CF0DEE6-E009-4EAE-9723-FC456357CBFC http://bioguid.osu.edu/xbiod_concepts/452212 Figures 3.2C, 3.24A–3.24F

Description. Body length of female: 8.78–9.20 mm (n=6). Body length of male: 7.60–7.83 mm (n=4). Color of A1: yellow to orange. A6 in female: distinctly longer than wide. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 2. Sculpture of dorsal A1: smooth. Color of head: black. Sculpture of frons directly above interantennal process: punctate rugose. Central keel: present, interrupted medially. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: dorsoventrally strigose. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: densely punctate. Sculpture of lateral pronotal area above pronotal

263 cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: absent. Macrosculpture of mesoscutal midlobe: sparsely punctate. Macrosculpture of lateral lobe of mesoscutum: punctate rugose. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: punctate rugose. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: present. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose anteriorly, smooth posteriorly. Color of legs: orange yellow to brown, with tarsi darker. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: sparse. Transverse sulcus on T2: present. Sculpture of T2: longitudinally punctate rugose. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: smooth to coriaceous. Posterior margin of T7 in male: emarginate medially between rounded projections. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. longitarsis, but it can be distinguished by the combination of the following characters: female A12 has two basiconic sensilla, and occipital carina complete. Etymology. The epithet is inspired by the Latin word for huge, in reference to the large body size of this species, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452212]

264 Material Examined. Holotype, female: ECUADOR: Orellana Prov., Yasuní Scientific Research Station (YSRS), 220m, 00.67°S 76.39°W, Yasuní National Park, 18.V–27.V.1990, P. Hibbs, OSUC 577500 (deposited in CNCI). Paratypes: (5 females, 4 males) BRAZIL: 4 females, 4 males, OSUC 149636, 202473–202478 (AEIC); OSUC 583458 (CNCI). ECUADOR: 1 female, OSUC 577499 (CNCI).

Chromoteleia levitas Chen & Johnson, sp. n. http://zoobank.org/F5087C96-A2E3-420E-B28F-7089D61A20A4 http://bioguid.osu.edu/xbiod_concepts/452159 Figures 3.25A–3.25F

Description. Body length of female: 5.68–6.60 mm (n=20). Body length of male: 4.60–5.10 mm (n=8). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: transversely striate to rugose. Central keel: present only in ventral portion of frons. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: punctate rugose dorsally and ventrally, strigose medially. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: punctate rugose anteriorly, smooth posteriorly. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose throughout. Macrosculpture of lateral lobe of mesoscutum: punctate rugose. Sculpture of notaulus: smooth. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: densely punctate rugose. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: largely smooth, punctate rugose anteriorly and

265 directly below femoral depression. Sculpture of dorsal metapleural area: rugose; smooth. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose throughout; rugose anteriorly, smooth posteriorly. Color of legs: orange yellow throughout. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: approximately as long as stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: sparse. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: densely punctate. Posterior margin of T7 in male: emarginate medially between rounded projections. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. rara, but it can be distinguished by the combination of the smooth notaulus smooth and striate A1. Etymology. The epithet is inspired by the Latin word for smoothness, in reference to the mostly smooth metapleuron, and is intended to be treated as a noun. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452159] Material Examined. Holotype, female: BRAZIL: RO, 62km S Ariquemes, Rancho Grande Farm, 12.XI–22.XI.1991, E. M. Fisher, OSUC 577476 (deposited in CNCI). Paratypes: (43 females, 8 males) BRAZIL: 21 females, 5 males, OSUC 149642, 202510 (AEIC); OSUC 577471–577474, 577477, 577479–577482, 584157–584158, 584331–584333 (CNCI); OSUC 354579, 380243 (MPEG); OSUC 577475, 577478, 577483, 584153, 584326 (OSUC); OSUC 225307–225308, 225311 (USNM). COLOMBIA: 5 females, OSUC 143982, 193662 (IAVH); OSUC 228672, 269686, 274926 (OSUC). COSTA RICA: 1 female, OSUC 584243 (CNCI). ECUADOR: 7 females, 2 males, OSUC 585091–585096, 585098, 586529 (CNCI); OSUC 586530 (OSUC). FRENCH GUIANA: 6 females, OSUC 555802, 555817, 555820, 555822, 586853 (CNCI); OSUC 555823 (OSUC). PERU: 2 females, 1 male, OSUC 584306, 584317

266 (CNCI); OSUC 320746 (USNM). SURINAME: 1 female, OSUC 584296 (CNCI). Other material: PERU: 1 female, 1 male, SEMC0983802, SEMC0985301T (KUNH).

Chromoteleia longitarsis Kieffer http://zoobank.org/17954F24-A488-4D04-99FE-12F22370E2C1 http://bioguid.osu.edu/xbiod_concepts/4214 Figures 3.5A–3.5B, 3.6F, 3.26A–3.27F, 3.42A

Chromoteleia longitarsis Kieffer, 1910a: 313, 314 (original description, keyed); Masner, 1976: 25 (type information); Johnson, 1992: 364 (cataloged, type information). Petalosema longitarsis (Kieffer): Kieffer, 1926: 358, 359 (generic transfer, description, keyed).

Description. Body length of female: 6.88–7.20 mm (n=20). Body length of male: 6.30–6.73 mm (n=20). Color of A1: yellow to orange. A6 in female: distinctly longer than wide. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: transversely striate to rugose. Central keel: complete, extending from interantennal process to median ocellus. Ventral margin of clypeus: straight. Granulate microsculpture of dorsal frons: absent. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: dorsoventrally strigose. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: sparsely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: rugose. Microsculpture of mesoscutum: absent. Macrosculpture of mesoscutal midlobe: punctate rugose throughout. Macrosculpture of lateral lobe of mesoscutum: densely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: densely punctate rugose. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: smooth. Mesopleural carina: present. Sculpture of mesepisternum below femoral

267 depression: punctate rugose. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: present. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: pale yellow with tarsi dark brown to black. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: sparse. Transverse sulcus on T2: absent. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: sinuate. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: densely punctate. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species can be distinguished from other Chromoteleia species by the following combination of characters: female A7 without basiconic sensilla, dorsal metapleural area with setae, basitarsus distinctly longer than remaining segments combined, postmarginal vein distinctly longer than stigmal vein, male T7 with posterior margin straight. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=4214] Material Examined. Holotype, male: BRAZIL: PA, no date, Baker, CAS TYPE9682 (deposited in CAS). Other material: (84 females, 58 males) BRAZIL: 7 females, 6 males, OSUC 202525, 202528, 202531 (AEIC); OSUC 584895–584896, 584898–584901 (CNCI); OSUC 379215 (MPEG); OSUC 185791, 56242, 584897 (OSUC). COLOMBIA: 2 females, OSUC 193428 (IAVH); OSUC 253477 (OSUC). COSTA RICA: 57 females, 26 males, OSUC 583420, 583776, 584260, 584263, 584857–584864, 584866–584868, 584870–584874, 584876– 584878, 584880–584887, 584889–584891, 584964–584969, 584971–584976, 584978–584980, 584982–584983, 584985–584986, 584988–584992, 584994–584999, 586751–586752, 586754, 586756, 586838, 586843 (CNCI); OSUC 584865, 584869, 584875, 584879, 584888, 584970, 584977, 584981, 584984, 584987, 584993, 585000, 586753 (OSUC). ECUADOR: 8 females, 1 male, OSUC 584892–584894, 584902, 584911–584912, 584919, 586528 (CNCI); DPI_FSCA

268 00010210 (CSCA). FRENCH GUIANA: 7 females, 11 males, OSUC 555803, 555806, 555808, 555810, 555812, 555816, 555818, 555824, 584913, 586239, 586424–586425, 586437, 586861 (CNCI); OSUC 555805, 555809, 555815, 586860 (OSUC). PANAMA: 2 males, OSUC 584380, 586755 (CNCI). PERU: 1 female, OSUC 578056 (CNCI). SURINAME: 1 female, OSUC 584903 (CNCI). TRINIDAD AND TOBAGO: 1 female, 12 males, OSUC 584144, 584904– 584910, 584914–584918 (CNCI).

Chromoteleia longa Chen & Johnson, sp. n. http://zoobank.org/1DB03949-CD59-4BC2-AD82-1DCFFA974FEB http://bioguid.osu.edu/xbiod_concepts/318200 Figures 3.1E–3.1F, 3.28A–2.28F

Description. Body length of female: 5.28–5.90 mm (n=20). Body length of male: 5.10–5.83 mm (n=20). Color of A1: yellow to orange. A6 in female: distinctly longer than wide. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: punctate rugose. Central keel: present, interrupted medially. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: narrowly smooth ventrally, punctate rugose dorsally. Color of mesosoma: orange. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: sparsely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: absent. Macrosculpture of mesoscutal midlobe: punctate rugose throughout. Macrosculpture of lateral lobe of mesoscutum: punctate rugose. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: densely punctate rugose. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral

269 depression: striate to densely punctate below mesopleural carina, otherwise smooth. Sculpture of dorsal metapleural area: rugose; smooth. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose anteriorly, smooth posteriorly. Color of legs: orange yellow throughout. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: black. Horn on T1 in female: absent. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: longitudinally punctate rugose. Sculpture of T6 in female: densely punctate and granulate. Length of T6 in female: approximately as long as wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: coriaceous anteriorly, densely punctate posteriorly. Posterior margin of T7 in male: emarginate medially between rounded projections. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. tricarinata, but it can be distinguished by the combination of the following characters: female A6 distinctly longer than wide, female A12 with one basiconic sensillum, median mesoscutal carina present anteriorly. Etymology. The epithet is inspired by the Latin word for long, in reference to the elongate antenna in female, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=318200] Material Examined. Holotype, female: COLOMBIA: Chocó Dept., visitor's center, M.815, 2m, 06°01'N 77°20'W, Utría Natural National Park, 28.IX–15.X.2000, Malaise trap, J. Pérez, OSUC 188702 (deposited in OSUC). Paratypes: (49 females, 59 males) BELIZE: 14 females, OSUC 556959, 556961, 556963–556964, 556967, 556969, 586771, 586869 (CNCI); OSUC 556958, 556960, 556962, 556965, 556968, 586867 (OSUC). COLOMBIA: 5 females, 11 males, OSUC 199631–199632 (FSCA); OSUC 144297, 190807–190808, 190819–190821 (IAVH); OSUC 144296, 144298, 188703, 190809, 190817–190818, 190822, 269594 (OSUC). COSTA RICA: 19 females, 39 males, OSUC 577985, 578006, 578008, 578071, 583424–583428, 583430–583433, 583435, 583438, 583442, 583445–583447, 583450–583452, 583454–583457, 583716, 583765, 583784–583785, 583799–583813, 583816, 583976, 584229, 584252, 586392,

270 586450 (CNCI); OSUC 583439, 583441, 583443–583444, 583453, 583814–583815 (OSUC). FRENCH GUIANA: 1 male, OSUC 586438 (CNCI). PANAMA: 11 females, 8 males, OSUC 556948–556949, 556956, 584178, 584374, 584377, 586176, 586376, 586622, 586677–586678, 586818 (CNCI); OSUC 320583, 556952, 586621 (OSUC); OSUC 221924, 320584, 320641, 320643 (TAMU).

Chromoteleia maura Chen & Masner, sp. n. http://zoobank.org/0C29F53A-F881-4B25-9AA7-D2ABB704F172 http://bioguid.osu.edu/xbiod_concepts/452214 Figures 3.29A–3.29F

Description. Body length of female: 5.20 mm (n=1). Color of A1: black. A6 in female: distinctly longer than wide. A5 in female: distinctly longer than wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: punctate rugose. Central keel: present, interrupted medially. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: dorsoventrally strigose. Color of mesosoma: black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: absent. Macrosculpture of mesoscutal midlobe: densely punctate. Macrosculpture of lateral lobe of mesoscutum: densely punctate. Sculpture of notaulus: smooth. Median mesoscutal carina: present along full length of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: densely punctate rugose. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: rugose anteriorly, sparsely punctate posteriorly. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: present. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose throughout. Color of legs: coxae and trochanters

271 orange yellow, remainder of the legs dark brown. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures. Length of postmarginal vein: distinctly shorter than stigmal vein. Color of metasoma in female: black. Horn on T1 in female: absent. Striae of posterior margin of T1 in female: dense. Transverse sulcus on T2: absent. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: approximately as long as wide. Shape of T6 in female in lateral view: sinuate. Apical spine on female T6: present. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species can be easily distinguished by its black body, mostly dark brown legs, and smooth notaulus. Etymology. The epithet is inspired by the Latin word for dark, in reference to the black body and legs, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452214] Material Examined. Holotype, female: ECUADOR: Napo Prov., 5km S Baeza, 1700m, 13.II.1983, Masner & Sharkey, OSUC 584757 (deposited in CNCI).

Chromoteleia parvitas Chen & Johnson, sp. n. http://zoobank.org/0EC98684-75B3-4609-BB53-9C64BC234AC9 http://bioguid.osu.edu/xbiod_concepts/452227 Figures 3.1C–3.1D, 3.30A–3.30F

Description. Body length of female: 3.38–3.65 mm (n=5). Color of A1: yellow to orange. A6 in female: distinctly wider than long. A5 in female: as long as wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: smooth. Color of head: black. Sculpture of frons directly above interantennal process: areolate. Central keel: complete, extending from interantennal process to median ocellus. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth medially, striate laterally. Sculpture of gena: coarsely punctate rugose.

272 Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly, sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: flat. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Sculpture of mesepisternum below femoral depression: punctate rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow throughout. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly shorter than stigmal vein. Color of metasoma in female: black. Horn on T1 in female: absent. Striae of posterior margin of T1 in female: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: approximately as long as wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. curta, but it can be distinguished by the combination of the following characters: A5 in female as long as wide, postmarginal vein distinctly shorter than stigmal vein, female T1 horn absent. Etymology. The epithet is inspired by the Latin word for small, in reference to the small body size of this species, and is intended to be treated as a noun. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452227] Material Examined. Holotype, female: ECUADOR: Sucumbíos Prov., Limoncocha, 250m, 15.VI–28.VI.1976, S. Peck & J. Peck, OSUC 586341 (deposited in CNCI). Paratypes: (4 females)

273 BRAZIL: 1 female, OSUC 584163 (CNCI). COLOMBIA: 2 females, OSUC 584303 (IAVH); OSUC 276182 (OSUC). ECUADOR: 1 female, OSUC 583752 (CNCI).

Chromoteleia pilus Chen & Johnson, sp. n. http://zoobank.org/6A3CC03A-FB31-4C4B-8BD3-87C2F935D18C http://bioguid.osu.edu/xbiod_concepts/452225 Figures 3.6C, 3.31A–3.31F

Description. Body length of female: 4.98–5.90 mm (n=20). Body length of male: 4.73–5.33 mm (n=20). Color of A1: yellow to orange. A6 in female: distinctly wider than long. A5 in female: distinctly longer than wide. A6 in male: as long as wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 2. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: punctate rugose. Central keel: present, interrupted medially. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: present. Occipital carina: interrupted medially. Granulate microsculpture of vertex: present. Sculpture of occiput: rugose. Sculpture of gena: narrowly smooth ventrally, punctate rugose dorsally. Color of mesosoma: orange. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: densely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose throughout. Macrosculpture of lateral lobe of mesoscutum: punctate rugose. Sculpture of notaulus: foveate. Notaular foveae: discrete. Median mesoscutal carina: present along full length of mesoscutum; present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: punctate throughout. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: present. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow

274 throughout. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: approximately as long as wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: coriaceous anteriorly, densely punctate posteriorly. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. depilis in color and size, but it can be distinguished by the combination of the following characters: dorsal metapleural area with setation, A6 in female as wide as long. Etymology. The epithet is inspired by the Latin word for hair, in reference to the setose dorsal metapleuron, and is intended to be treated as a noun. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452225] Material Examined. Holotype, female: COSTA RICA: Puntarenas Prov., road to Rincón, 24km W Pan-American Highway, 200m, II-1989 – III-1989, Hanson & Gauld, OSUC 584261 (deposited in CNCI). Paratypes: (53 females, 31 males) COSTA RICA: 49 females, 31 males, OSUC 149651, 202550, 202553, 202555–202556, 203109 (AEIC); OSUC 556972, 556986, 556988, 556992, 557002, 557011–557012, 557049, 557061, 557083, 577903, 577907, 577933, 577935–577937, 577941, 577953, 577988, 578002–578004, 578058, 578082–578083, 578103, 583440, 583448, 583717, 583723, 583817, 583880, 583887, 583943–583944, 584032, 584035, 584037–584039, 584236, 584238, 584262, 586170, 586173, 586210, 586290, 586310, 586512, 586533, 586544, 586644, 586680, 586747, 586772, 586836 (CNCI); OSUC 202551, 202554, 556989, 557040, 56234, 578005, 578081, 578091, 583882, 583886, 583940, 584041, 584247, 586509, 586545, 586646, 586780 (OSUC); OSUC 221941 (TAMU). MEXICO: 1 female, OSUC 584145 (CNCI). PANAMA: 3 females, OSUC 584378, 584381, 586504 (CNCI).

Chromoteleia plana Chen & Johnson, sp. n.

275 http://zoobank.org/DA401CB2-DCD9-405A-83AE-8159593A1510 http://bioguid.osu.edu/xbiod_concepts/452218 Figures 3.32A–3.32F

Description. Body length of female: 7.31 mm (n=1). Color of A1: yellow to orange. A6 in female: distinctly wider than long. A5 in female: as long as wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate; smooth. Color of head: black. Sculpture of frons directly above interantennal process: areolate. Central keel: present only in ventral portion of frons. Ventral margin of clypeus: straight. Granulate microsculpture of dorsal frons: absent. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: narrowly smooth ventrally, punctate rugose dorsally. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: sparsely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: absent. Macrosculpture of mesoscutal midlobe: sparsely punctate at anterior margin, smooth posteriorly. Macrosculpture of lateral lobe of mesoscutum: smooth. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Shape of mesoscutellum: strongly transverse. Mesoscutellum in lateral view: flat. Sculpture of mesoscutellum: smooth throughout. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: areolate. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: largely smooth, punctate rugose anteriorly and directly below femoral depression. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow with coxae dark brown. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: approximately as long as stigmal vein.

276 Color of metasoma in female: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices, with a narrow smooth strip medially. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of medial S2: punctate rugose, with a narrow smooth strip medially. Diagnosis. This species can be easily distinguished by the following characters: body extremely elongate, central keel present only in ventral portion of frons, mesoscutellum strongly transverse, and T2–T6 with with a narrow smooth strip medially. Etymology. The epithet is inspired by the Latin word for flat or plain, in reference to the flat and smooth mesoscutellum, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452218] Material Examined. Holotype, female: FRENCH GUIANA: Cayenne Arrond., 323m, 04°33.998'N 52°12.442'W, Kaw Mountains, 15.XII–18.XII.2011, Malaise trap, A. Desjardins, OSUC 577460 (deposited in CNCI).

Chromoteleia rara Chen & Johnson, sp. n. http://zoobank.org/0E7CB667-807C-4C4B-9858-BFE942BBAD19 http://bioguid.osu.edu/xbiod_concepts/452222 Figures 3.2A–3.2B, 3.33A–3.33F

Description. Body length of female: 4.68–5.30 mm (n=20). Body length of male: 4.50–5.13 mm (n=20). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: as long as wide. Number of basiconic sensilla on A7: 1; 2. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: smooth. Color of head: black. Sculpture of frons directly above interantennal process: areolate. Central keel: present only in ventral portion of frons. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: dorsoventrally strigose.

277 Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly, sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: densely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Sculpture of mesepisternum below femoral depression: punctate rugose. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow throughout. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: distinctly shorter than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T7 in male: coriaceous anteriorly, densely punctate posteriorly. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. cuneus, with which it shares a short postmarginal vein, and it can be distinguished by the rounded apex of T6 in dorsal view. Etymology. The epithet is inspired by the Latin word for uncommon. We apply it to this species because this species is the only species in Chromoteleia that can have either one or two basiconic sensilla on A7. The name is treated as an adjective in apposition. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452222]

278 Material Examined. Holotype, female: COSTA RICA: Heredia Prov., Puerto Viejo, La Selva Biological Station, 100m, II.1993 – III.1993, Malaise trap, P. Hanson, OSUC 577495 (deposited in CNCI). Paratypes: (45 females, 28 males) BELIZE: 2 females, 1 male, OSUC 577509– 577510 (CNCI); OSUC 93700 (OSUC). BRAZIL: 1 male, OSUC 586574 (CNCI). COLOMBIA: 3 females, OSUC 199606, 199612 (FSCA); OSUC 268912 (OSUC). COSTA RICA: 27 females, 17 males, OSUC 202552 (AEIC); OSUC 253964, 556978–556979, 556981, 557005–557006, 577489, 577491, 577493, 577904, 577906, 577945–577946, 577973, 577979, 577991, 578079, 578086, 583758, 583879, 583898, 583925, 583949, 584025, 584235, 586267, 586378, 586480, 586618, 586662, 586664 (CNCI); OSUC 556997, 577490, 577492, 577494, 577912, 577934, 577939, 577948, 584010, 586782, 586784 (OSUC); OSUC 204983 (UCDC). ECUADOR: 3 females, 8 males, OSUC 202545–202548 (AEIC); OSUC 577432, 577434– 577435, 584106, 584756 (CNCI); OSUC 199608, 199610 (FSCA). FRENCH GUIANA: 1 female, OSUC 47022 (OSUC). HONDURAS: 2 females, OSUC 369621, 413783 (MZLU). PANAMA: 6 females, 1 male, OSUC 586819 (CNCI); OSUC 320639 (OSUC); OSUC 221918– 221920, 320644 (TAMU); OSUC 205000 (UCDC). TRINIDAD AND TOBAGO: 1 female, OSUC 586288 (CNCI). Comments. This species is extremely variable in body length, which may be an indication that it attacks multiple species with different size eggs, or that the eggs of its host(s) vary widely in size.

Chromoteleia robusta Chen & Johnson, sp. n. http://zoobank.org/A38A572D-E20C-418F-8988-235126D46E8E http://bioguid.osu.edu/xbiod_concepts/452215 Figures 3.3B, 3.35A–3.35F

Description. Body length of female: 5.18–5.53 mm (n=7). Body length of male: 4.30–4.93 mm (n=2). Color of A1: yellow to orange. A6 in female: distinctly longer than wide. A5 in female: distinctly longer than wide. A6 in male: as long as wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 2. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: punctate rugose. Central keel: complete, extending from interantennal process to median ocellus. Ventral margin of clypeus: straight.

279 Granulate microsculpture of dorsal frons: absent. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth medially, striate laterally. Sculpture of gena: dorsoventrally strigose. Color of mesosoma: variably orange to black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose throughout. Macrosculpture of lateral lobe of mesoscutum: punctate rugose. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: absent. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: densely punctate rugose. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: punctate rugose. Sculpture of dorsal metapleural area: rugose; smooth. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose anteriorly, smooth posteriorly. Color of legs: orange yellow throughout. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: densely punctate. Length of postmarginal vein: approximately as long as stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: absent. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: approximately as long as wide; wider than long. Shape of T6 in female in lateral view: sinuate. Apical spine on female T6: present. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: granulate. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Length of postmarginal vein: approximately as long as stigmal vein. Diagnosis. This species is similar to C. feng in the acute apex of T6, and it can be distinguished by the combination of the following characters: female A12 with two basiconic sensilla, occipital carina interrupted medially, notaulus foveate.

280 Etymology. The epithet is inspired by the Latin word for strong or robust, in reference to the robust appearance of this species, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452215] Material Examined. Holotype, female: VENEZUELA: Aragua St., La Cumbre trail, Rancho Grande, IV.1987 – V.1987, flight intercept trap, OSUC 577466 (deposited in CNCI). Paratypes: VENEZUELA: 6 females, 2 males, OSUC 210349 (AMNH); OSUC 577462–577465, 577467– 577469 (CNCI).

Chromoteleia rufithorax Kieffer http://zoobank.org/ECD55F87-65C5-4C1E-93DC-BB6974CA164A http://bioguid.osu.edu/xbiod_concepts/4215

Chromoteleia rufithorax Kieffer, 1907: 266 (original description, keyed); Kieffer, 1908a: 25 (described as new); Kieffer, 1910a: 313 (keyed); Johnson, 1992: 364 (cataloged). Petalosema rufithorax (Kieffer): Kieffer, 1926: 358 (generic transfer, description, keyed).

Comments. We were not able to locate the type specimens of this species, and its status and identity are unclear.

Chromoteleia semicyanea Ashmead http://zoobank.org/5D560202-AF14-47AB-AB0A-70E96B8665C7 http://bioguid.osu.edu/xbiod_concepts/4216 Figures 3.36A–3.36F

Chromoteleia semicyanea Ashmead, 1893: 220 (original description); Ashmead, 1894: 224 (redescribed as new); Ashmead, 1900: 327 (distribution); Kieffer, 1907: 266 (keyed); Kieffer, 1910a: 312 (keyed); Kieffer, 1926: 406 (description); Masner, 1976: 25 (type information, lectotype designation); Johnson, 1992: 364 (cataloged, type information).

Description. Body length of female: 5.48–5.60 mm (n=2). Body length of male: 4.80–4.93 mm (n=3). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly

281 longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate; smooth. Color of head: blue. Sculpture of frons directly above interantennal process: transversely striate to rugose. Central keel: absent. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: dorsoventrally strigose. Color of mesosoma: blue dorsally, black laterally. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: densely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: rugose. Microsculpture of mesoscutum: coriaceous. Macrosculpture of mesoscutal midlobe: sparsely punctate. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: absent. Sculpture of mesepisternum below femoral depression: punctate throughout. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow throughout. Length of hind basitarsus: distinctly longer than remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: orange. Color of metasoma in male: orange. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely

282 punctate. Sculpture of T7 in male: smooth anteriorly, rugulose posteriorly. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely punctate to punctate rugose. Diagnosis. This species can be distinguished from other Chromoteleia species by the following combination of characters: head and dorsal mesosoma blue, metasoma orange, frons without central keel, mesopleural carina absent. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=4216] Material Examined. Lectotype, male: SAINT VINCENT AND THE GRENADINES: Saint Vincent Island, 2000ft, no date, H. H. Smith, USNM Type No. 2251 (deposited in USNM). Other material: SAINT VINCENT AND THE GRENADINES: 2 females, 2 males, OSUC 584763–584766 (CNCI).

Chromoteleia semilutea Chen & Johnson, sp. n. http://zoobank.org/7F8871F3-3D8E-4985-9CC5-850F70A1558F http://bioguid.osu.edu/xbiod_concepts/452219 Figures 3.37A–3.37F

Description. Body length of female: 5.28–5.50 mm (n=10). Body length of male: 4.30–4.83 mm (n=3). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: as long as wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 2. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: transversely striate to rugose. Central keel: complete, extending from interantennal process to median ocellus. Ventral margin of clypeus: straight. Granulate microsculpture of dorsal frons: absent. Occipital carina: interrupted medially. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth. Sculpture of gena: dorsoventrally strigose. Color of mesosoma: orange. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose throughout. Macrosculpture of lateral lobe of mesoscutum: punctate rugose. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present along full length of mesoscutum.

283 Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: densely punctate rugose. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: smooth. Mesopleural carina: absent. Sculpture of mesepisternum below femoral depression: largely smooth, punctate rugose anteriorly and directly below femoral depression. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow throughout. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: mostly black with T1–T3 orange to dark brown. Color of metasoma in male: mostly black with T1–T3 orange to dark brow. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: approximately as long as wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: granulate. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely punctate to punctate rugose. Diagnosis. This species can be distinguished from other Chromoteleia species by the following combination of characters: metasoma with T1–T3 orange to dark brown, frons with central keel complete, mesopleural carina absent. Etymology. The name semilutea refers to the half orange metasoma of this species and is used as an adjective in apposition. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452219] Material Examined. Holotype, female: BELIZE: Orange Walk Dist., Rio Bravo Conservation and Management Area, 15.VII–21.VII.1996, Malaise trap, P. Kovarik, OSUC 577506 (deposited in CNCI). Paratypes: (9 females, 3 males) BELIZE: 3 females, 1 male, OSUC 577505 (CNCI); OSUC 225288, 232143, 47912 (OSUC). MEXICO: 6 females, 2 males, OSUC 556945–556946, 577501–577504, 586774 (CNCI); OSUC 586773 (OSUC).

284

Chromoteleia sparsa Chen & Johnson, sp. n. http://zoobank.org/C074AD5B-E19B-4F8F-B403-BE9B5A81AEA5 http://bioguid.osu.edu/xbiod_concepts/452213 Figures 3.38A–3.39F

Description. Body length of female: 5.38–5.70 mm (n=20). Body length of male: 4.90–5.15mm (n=20). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 1. Number of basiconic sensilla on A12: 1. Sculpture of dorsal A1: punctate; smooth. Color of head: black. Sculpture of frons directly above interantennal process: areolate. Central keel: present, interrupted medially. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: absent. Occipital carina: complete. Granulate microsculpture of vertex: absent. Sculpture of occiput: smooth medially, striate laterally. Sculpture of gena: punctate rugose dorsally and ventrally, strigose medially. Color of mesosoma: variably orange to nearly black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: densely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: rugose. Microsculpture of mesoscutum: absent. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly, sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: flat. Sculpture of mesoscutellum: smooth medially, densely punctate laterally. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: absent or indistinguishable from sculpture. Sculpture of metascutellum: areolate. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: punctate throughout. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: absent. Setation of area directly dorsal to the metapleural triangle: present. Sculpture of ventral metapleural area: rugose throughout. Color of legs: orange yellow throughout. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures.

285 Length of postmarginal vein: distinctly shorter than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: sparse. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5x longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: granulate. Posterior margin of T7 in male: emarginate medially between rounded projections. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species is similar to C. copiosa, but it can be distinguished by the combinations of the following characters: occipital carina complete, netrion rugose, and the hind basitarsus approximately as long as the remaining segments combined. Etymology. The epithet is inspired by the Latin word for scattered, in reference to the sparsely punctate posterior part of mesoscutum, and is intended to be treated as an adjective. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=452213] Material Examined. Holotype, female: ECUADOR: Sucumbíos Prov., 270m, 00°30'S 76°30'W, Sacha Lodge, 13.VI–23.VI.1994, Malaise trap, P. Hibbs, OSUC 584751 (deposited in CNCI). Paratypes: (64 females, 39 males) ARGENTINA: 1 female, OSUC 584143 (CNCI). BRAZIL: 36 females, 30 males, OSUC 149640–149641, 202482, 202504–202505, 202512, 202514, 202536 (AEIC); OSUC 557091, 583487–583489, 583493, 583499, 583502, 583504, 583509, 584149, 586149, 586259, 586343–586344, 586347–586349, 586353–586355, 586361, 586577–586581, 586586–586587, 586596, 586598, 586600, 586603–586608 (CNCI); OSUC 199635 (FSCA); OSUC 225586, 241253 (MPEG); OSUC 147541, 246558, 254870 (MZSP); OSUC 138674, 147540, 225227, 235205, 246557, 254868–254869, 354648, 583486, 583500, 586350–586351, 586585, 586590 (OSUC); OSUC 204990 (UCDC). COLOMBIA: 2 females, 2 males, OSUC 584302 (CNCI); OSUC 182714, 230409 (IAVH); OSUC 230410 (OSUC). COSTA RICA: 4 females, 1 male, OSUC 577923–577924, 586498, 586641 (CNCI); OSUC 586175 (OSUC). ECUADOR: 9 females, 2 males, OSUC 556944, 584748–584750, 584753– 584755, 586339–586340 (CNCI); OSUC 583707, 584752 (OSUC). FRENCH GUIANA: 7 females, OSUC 555807, 555813, 586435, 586448, 586459 (CNCI); OSUC 555811, 586863

286 (OSUC). GUYANA: 1 female, OSUC 583434 (CNCI). MEXICO: 2 males, OSUC 584295 (CNCI); OSUC 221932 (TAMU). TRINIDAD AND TOBAGO: 1 male, OSUC 586308 (CNCI). VENEZUELA: 4 females, 1 male, OSUC 557088, 584141, 586362 (CNCI); OSUC 251644, 251662 (MIZA). Comments. This species is extremely variable in mesosoma color and can be roughly divided into dark and light forms.

Chromoteleia tricarinata Kieffer http://zoobank.org/301D1A78-C6C1-4A91-8638-D3E7237C7611 http://bioguid.osu.edu/xbiod_concepts/4218 Figures 3.1A–3.1B, 3.40A–3.42A

Chromoteleia rufithorax var. tricarinata Kieffer, 1909: 250 (original description). Petalosema tricarinata (Kieffer): Kieffer, 1926: 358, 359 (generic transfer, description, change to species status, keyed). Chromoteleia tricarinata Kieffer: Masner, 1976: 25 (type information, taxonomic status); Johnson, 1992: 364 (cataloged, type information).

Description. Body length of female: 5.28–5.90 mm (n=20). Body length of male: 5.20–5.83 mm (n=20). Color of A1: yellow to orange. A6 in female: as wide as long. A5 in female: distinctly longer than wide. A6 in male: approximately 2.0x longer than wide. Number of basiconic sensilla on A7: 0. Number of basiconic sensilla on A12: 2. Sculpture of dorsal A1: striate. Color of head: black. Sculpture of frons directly above interantennal process: punctate rugose. Central keel: complete, extending from interantennal process to median ocellus. Ventral margin of clypeus: pointed. Granulate microsculpture of dorsal frons: present. Occipital carina: interrupted medially. Granulate microsculpture of vertex: present. Sculpture of occiput: smooth medially, striate laterally. Sculpture of gena: coarsely punctate rugose. Color of mesosoma: orange. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: smooth. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth throughout. Sculpture of netrion: transversely striate. Microsculpture of mesoscutum: granulate. Macrosculpture of mesoscutal midlobe: punctate rugose throughout. Macrosculpture of lateral

287 lobe of mesoscutum: punctate rugose. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: absent. Mesoscutellum in lateral view: convex. Sculpture of mesoscutellum: densely punctate rugose. Shape of metascutellum: trapezoidal with broad posterior margin. Median metascutellar carina: present. Sculpture of metascutellum: rugose. Sculpture of lateral propodeal area: rugose. Mesopleural carina: present. Sculpture of mesepisternum below femoral depression: punctate rugose. Sculpture of dorsal metapleural area: rugose. Setation of dorsal metapleural area: present. Setation of area directly dorsal to the metapleural triangle: absent. Sculpture of ventral metapleural area: rugose anteriorly, obliquely striate posteriorly. Color of legs: orange yellow throughout. Length of hind basitarsus: about as long as remaining segments combined. Sculpture of hind coxa: largely smooth, with sparse fine punctures. Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: approximately as long as wide; wider than long. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: granulate. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices. Diagnosis. This species can be distinguished from other Chromoteleia by the following combination of characters: female A7 without basiconic sensillum, mesoscutum without median mesoscutal carina, dorsal metapleural area with setae, ventral metapleural area obliquely striate, male T7 with posterior margin straight. Link to Distribution Map. [http://hol.osu.edu/map-large.html?id=4218] Material Examined. Holotype, male, C. rufithorax var. tricarinata: BELIZE: no date, CAS TYPE9759 (deposited in CAS). Other material: (330 females, 551 males) BELIZE: 32 females, 27 males, OSUC 556957, 584778–584798, 584800–584804, 584806, 584921, 584932–584934, 586445, 586758, 586870–586871 (CNCI); OSUC 186068, 186070, 225287, 232140–232142, 47918–47919, 47940, 48011, 48037, 64021, 64026–64028, 64046–64047, 93534, 93586–93588, 94076–94077 (OSUC). BRAZIL: 1 female, 2 males, OSUC 56227–56229 (OSUC).

288 COLOMBIA: 1 female, OSUC 586287 (CNCI). COSTA RICA: 191 females, 340 males, OSUC 149653, 149657, 202538 (AEIC); OSUC 232992 (BPBM); OSUC 232058, 232120– 232138, 232188, 232190–232191, 374050, 556994, 556998, 557003, 557007, 557009, 557015, 557017, 557020–557022, 557024–557029, 557046–557047, 557052, 557054–557057, 557059– 557060, 557063–557080, 557082, 577894, 577897, 577900–577901, 577905, 577925, 577928, 577930–577931, 577942, 577952, 577956–577961, 577963, 577965–577970, 577972, 578010– 578011, 578019, 578023–578024, 578044, 578046, 578053, 578059, 578063–578065, 578067– 578070, 578072, 578074–578075, 578077–578078, 578084, 578087–578090, 578093–578096, 578098, 578100–578102, 578105, 583414, 583417–583418, 583421, 583429, 583437, 583460, 583462, 583465–583466, 583470, 583514–583515, 583518–583520, 583709–583715, 583718, 583720–583722, 583724–583730, 583732–583738, 583740–583750, 583754–583755, 583759, 583761–583762, 583766–583767, 583772, 583774–583775, 583777–583783, 583786, 583788– 583793, 583795, 583797–583798, 583818–583828, 583830–583833, 583835–583849, 583851– 583877, 583883–583885, 583888, 583893–583895, 583899–583900, 583902–583909, 583911– 583912, 583914–583915, 583917–583924, 583929–583930, 583939, 583941–583942, 583945– 583946, 583950–583951, 583977, 583979, 583983–583987, 583991, 583995–583996, 583999– 584002, 584007, 584013, 584015, 584018, 584020–584022, 584024, 584029–584030, 584034, 584036, 584040, 584043–584044, 584046, 584138, 584225, 584227, 584230, 584233, 584240, 584242, 584244, 584250, 584264–584270, 584274–584276, 584281–584283, 584287, 585121– 585124, 585126–585144, 586127–586128, 586139–586141, 586152–586156, 586165, 586172, 586174, 586184, 586195, 586197, 586199, 586202, 586204, 586207–586208, 586211, 586215, 586218–586221, 586262–586266, 586270, 586275–586286, 586291–586292, 586296–586301, 586374–586375, 586379–586389, 586391, 586405, 586407–586410, 586414, 586473, 586481– 586484, 586486, 586488, 586490–586492, 586495–586497, 586506, 586508, 586511, 586514– 586515, 586535–586537, 586540, 586560–586564, 586566, 586569–586571, 586628, 586630, 586632, 586634, 586636–586637, 586642–586643, 586647, 586650–586655, 586657, 586660– 586661, 586663, 586665–586666, 586671, 586763, 586765–586768, 586779, 586835, 586872 (CNCI); DPI_FSCA 00010211 (CSCA); SM0810075, SM0810348T (KUNH); OSUC 185844, 374051–374054 (OSUC); OSUC 320638, 320648–320650 (TAMU); OSUC 204982, 204999 (UCDC); OSUC 157812 (UCMC). GUATEMALA: 9 females, 16 males, OSUC 584920, 584922–584924, 584926–584931, 584935–584936, 584940–584948, 584950, 584952–584953

289 (CNCI); OSUC 317963 (OSUC). HONDURAS: 13 females, 16 males, OSUC 584961–584963 (CNCI); OSUC 369620, 369622, 369624, 369626–369627, 413758–413759, 413761–413763, 413766–413771, 413773–413782 (MZLU). MEXICO: 14 females, 20 males, OSUC 556947, 584937–584939, 584949, 584951, 584954–584960 (CNCI); OSUC 268812–268813, 268815– 268818, 271011, 271013–271014, 271016, 56230–56233, 56235, 56237–56241, 56243 (OSUC). NICARAGUA: 1 female, 1 male, DPI_FSCA 00010212 (CSCA); OSUC 204951 (UCDC). PANAMA: 68 females, 128 males, OSUC 149652, 149655, 202542–202543, 202557, 202559, 202561–202575 (AEIC); OSUC 557099–557112, 578054, 583459, 583483, 583501, 583505– 583507, 584179–584181, 584183–584224, 584337–584373, 584375–584376, 584383–584397, 586158, 586345, 586373, 586377, 586465–586468, 586477–586479, 586521–586527, 586565, 586613–586617, 586627, 586631, 586682–586692, 586815–586817, 586820–586823, 586825, 586874 (CNCI); OSUC 221922–221923, 319208–319209, 320642, 320645–320647, 321353 (TAMU). VENEZUELA: 1 male, OSUC 557086 (CNCI).

290

Figure 3.1 A–B, Chromoteleia tricarinata Kieffer, female (OSUC 268814). A, Antenna; B, Antennal clava, ventral view.C–D, Chromoteleia parvitas sp. n., holotype, female (OSUC 276182). C, Antenna; D, Antennal clava, ventral view; E–F, Chromoteleia longa sp. n., holotype, female (OSUC 320643). E, Antenna; F, Antennal clava, ventral view.

291

Figure 3.2 A–B, Chromoteleia rara sp. n., female, holotype (OSUC 577495). A, Antenna; B, Antennal clava, ventral view.C, Chromoteleia ingens sp. n., paratype, male (OSUC 583458), Antenna; D, Chromoteleia bidens sp. n., paratype, male (OSUC 185675), Antenna.

292

Figure 3.3 Wings A, Chromoteleia longitarsis Kieffer, female (OSUC 584896). B, Chromoteleia robusta sp. n., female, paratype (OSUC 577466). C, Chromoteleia cuneus sp. n., female, holotype (OSUC 585001).

293

Figure 3.4 Mesopleuron and metapleuron. A, Chromoteleia connectens Kieffer, female, holotype (CAS TYPE9618). B, Chromoteleia longitarsis Kieffer, female (OSUC 584896).

294

Figure 3.5 A–B, Chromoteleia longitarsis Kieffer, female (OSUC 584896). A, Apical metasoma, dorsal view; B, Apical metasoma, lateral view. C–D Chromoteleia depilis sp. n., female, holotype (OSUC 577436). C, Apical metasoma, dorsal view; D, Apical metasoma, lateral view. E–F, Chromoteleia bidens sp. n., female, holotype (OSUC 577455). E, Apical metasoma, dorsal view; F, Apical metasoma, lateral view.

295

Figure 3.6 A–B Chromoteleia fuscicornis Kieffer, female (OSUC 584322). A, Apical metasoma, dorsal view; B, Apical metasoma, lateral view. C, Chromoteleia pilus sp. n., male, paratype (OSUC 577941), Apical metasoma, dorsal view. D, Chromoteleia fuscicornis Kieffer, male (OSUC 586417), Apical metasoma, dorsal view. E, Chromoteleia bidens sp. n., male, paratype (OSUC 577455), Apical metasoma, dorsal view Apical. F, Chromoteleia longitarsis Kieffer, female (OSUC 584896), Hind tarsus. G, Chromoteleia depilis sp. n., female, paratype (OSUC 577436)), Hind tarsus. H, Chromoteleia bidens sp. n., female, paratype (OSUC 577455), Hind tarsus.

296

Figure 3.7 Chromoteleia aequalis sp. n., female, holotype (OSUC 221942). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

297

Figure 3.8 Chromoteleia alternata sp. n., female, holotype (OSUC 136573). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

298

Figure 3.9 Chromoteleia bidens sp. n., female, holotype (OSUC 577455). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

299

Figure 3.10 Oxyscelio congoana Risbec, male, holotype (EY11376). A, Lateral habitus; B, Dorsal habitus; C, Head, anterior view; D, Head and mesosoma, dorsal view.

300

Figure 3.11 Chromoteleia congoana (Risbec), female (OSUC 207829). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

301

Figure 3.12 Chromoteleia connectens Kieffer, female, holotype (CAS TYPE9618). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

302

Figure 3.13 Chromoteleia brevitarsis Kieffer, female, holotype (CAS TYPE9597). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

303

Figure 3.14 Chromoteleia copiosa sp. n., female, holotype (OSUC 577518). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

304

Figure 3.15 Chromoteleia cuneus sp. n., female, holotype (OSUC 585001). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

305

Figure 3.16 Chromoteleia curta sp. n., female, holotype (OSUC 185626). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: light form)

306

Figure 3.17 Chromoteleia curta sp. n., female, paratype (OSUC 149925). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: dark form)

307

Figure 3.18 Chromoteleia depilis sp. n., female, holotype (OSUC 577436). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

308

Figure 3.19 Chromoteleia dispar sp. n., female, holotype (OSUC 190774). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

309

Figure 3.20 Chromoteleia feng sp. n., female, holotype (OSUC 233344). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

310

Figure 3.21 Chromoteleia fossa sp. n., female, holotype (OSUC 320582). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

311

Figure 3.22 Chromoteleia fuscicornis Kieffer, female, holotype (CAS TYPE9649). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head, anterior view.

312

Figure 3.23 Chromoteleia fuscicornis Kieffer, female (OSUC 584322). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

313

Figure 3.24 Chromoteleia ingens sp. n., female, holotype (OSUC 577500). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

314

Figure 3.25 Chromoteleia levitas sp. n., female, holotype (OSUC 577476). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

315

Figure 3.26 Chromoteleia longitarsis Kieffer, male, holotype (CAS TYPE9682). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

316

Figure 3.27 Chromoteleia longitarsis Kieffer, female (OSUC 584896). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

317

Figure 3.28 Chromoteleia longa sp. n., female, holotype (OSUC 188702). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

318

Figure 3.29 Chromoteleia maura sp. n., female, holotype (OSUC 584757). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

319

Figure 3.30 Chromoteleia parvitas sp. n., female, holotype (OSUC 586341). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

320

Figure 3.31 Chromoteleia pilus sp. n., female, holotype (OSUC 584261). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

321

Figure 3.32 Chromoteleia plana sp. n., female, holotype (OSUC 577460). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

322

Figure 3.33 Chromoteleia rara sp. n., female, holotype (OSUC 577495). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: small form)

323

Figure 3.34 Chromoteleia rara sp. n., female, paratype (OSUC 557090). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: large form)

324

Figure 3.35 Chromoteleia robusta sp. n., female, holotype (OSUC 577466). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

325

Figure 3.36 Chromoteleia semicyanea Ashmead, male, holotype (USNM Type No. 2251). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, female (OSUC 584765), Metasoma, dorsal view.

326

Figure 3.37 Chromoteleia semilutea sp. n., female, holotype (OSUC 577506). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

327

Figure 3.38 Chromoteleia sparsa sp. n., female, holotype (OSUC 584751). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: light form)

328

Figure 3.39 Chromoteleia sparsa sp. n., female, paratype (OSUC 583488). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view. (NOTE: dark form)

329

Figure 3.40 Chromoteleia tricarinata Kieffer, male, holotype (CAS TYPE9759). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

330

Figure 3.41 Chromoteleia tricarinata Kieffer, female (OSUC 584805). A, Lateral habitus; B, Head and mesosoma, lateral view; C, Dorsal habitus; D, Head and mesosoma, dorsal view; E, Head, anterior view; F, Metasoma, dorsal view.

331

Figure 3.42 Mesoscutum, mmc: median mesoscutal carina. A, Chromoteleia tricarinata Kieffer, male, holotype (CAS TYPE9759). B, Chromoteleia connectens Kieffer, female, holotype (CAS TYPE9618). C, Chromoteleia longitarsis Kieffer, male, holotype (CAS TYPE9682). D, Chromoteleia curta sp. n., female, holotype (OSUC 185626).

332 Table 3.1 List of morphological terms and links to Hymenoptera Anatomy Ontology database for New World Calliscelio.

A1 http://purl.obolibrary.org/obo/HAO_0000908 A2 http://purl.obolibrary.org/obo/HAO_0000706 A3 http://purl.obolibrary.org/obo/HAO_0001148 A7 http://purl.obolibrary.org/obo/HAO_0001885 A12 http://purl.obolibrary.org/obo/HAO_0001884 antenna http://purl.obolibrary.org/obo/HAO_0000101 antennomere http://purl.obolibrary.org/obo/HAO_0000107 area http://purl.obolibrary.org/obo/HAO_0000146 body http://purl.obolibrary.org/obo/HAO_0000182 carina http://purl.obolibrary.org/obo/HAO_0000188 central keel http://purl.obolibrary.org/obo/HAO_0000109 cpa cervical pronotal area http://purl.obolibrary.org/obo/HAO_0000194 clava http://purl.obolibrary.org/obo/HAO_0000203 clypeus http://purl.obolibrary.org/obo/HAO_0000212 compound eye http://purl.obolibrary.org/obo/HAO_0000217 coxa http://purl.obolibrary.org/obo/HAO_0000228 depression http://purl.obolibrary.org/obo/HAO_0000241 dpa dorsal pronotal area http://purl.obolibrary.org/obo/HAO_0000267 egg http://purl.obolibrary.org/obo/HAO_0000286 epomial carina http://purl.obolibrary.org/obo/HAO_0000307 eye http://purl.obolibrary.org/obo/HAO_0000217 femur http://purl.obolibrary.org/obo/HAO_0000327 fore wing http://purl.obolibrary.org/obo/HAO_0000351 frons http://purl.obolibrary.org/obo/HAO_0001523 gena http://purl.obolibrary.org/obo/HAO_0000371 head http://purl.obolibrary.org/obo/HAO_0000397 hind coxa http://purl.obolibrary.org/obo/HAO_0000587 hind tibia http://purl.obolibrary.org/obo/HAO_0000631 hind wing http://purl.obolibrary.org/obo/HAO_0000400 inner orbit http://purl.obolibrary.org/obo/HAO_0000419 interantennal process http://purl.obolibrary.org/obo/HAO_0000422 lateral lobe of mesoscutum http://purl.obolibrary.org/obo/HAO_0000466 lateral ocellus http://purl.obolibrary.org/obo/HAO_0000481 LOL lateral ocellar line http://purl.obolibrary.org/obo/HAO_0000480 lpa lateral pronotal area http://purl.obolibrary.org/obo/HAO_0000483 malar sulcus http://purl.obolibrary.org/obo/HAO_0000504 mandible http://purl.obolibrary.org/obo/HAO_0000506 margin http://purl.obolibrary.org/obo/HAO_0000510 mesepisternum http://purl.obolibrary.org/obo/HAO_0001872 med mesopleural depression http://purl.obolibrary.org/obo/HAO_0000326 mesopleuron http://purl.obolibrary.org/obo/HAO_0000566

333 Table 3.1. Continued mesoscutellum http://purl.obolibrary.org/obo/HAO_0000574 mesoscutum http://purl.obolibrary.org/obo/HAO_0001490 mesosoma http://purl.obolibrary.org/obo/HAO_0000576 metapleuron http://purl.obolibrary.org/obo/HAO_0000621 metascutellum http://purl.obolibrary.org/obo/HAO_0000625 metasoma http://purl.obolibrary.org/obo/HAO_0000626 midlobe of mesoscutum http://purl.obolibrary.org/obo/HAO_0000520 netrion http://purl.obolibrary.org/obo/HAO_0000644 notauli (notaulus) http://purl.obolibrary.org/obo/HAO_0000647 occipital carina http://purl.obolibrary.org/obo/HAO_0000653 ocellus http://purl.obolibrary.org/obo/HAO_0000661 ot ocellar triangle http://purl.obolibrary.org/obo/HAO_0000430 OOL ocular ocellar line http://purl.obolibrary.org/obo/HAO_0000662 orbit http://purl.obolibrary.org/obo/HAO_0000672 POL posterior ocellar line http://purl.obolibrary.org/obo/HAO_0000759 process http://purl.obolibrary.org/obo/HAO_0000822 propodeum http://purl.obolibrary.org/obo/HAO_0001248 S1 http://purl.obolibrary.org/obo/HAO_0001997 S2 http://purl.obolibrary.org/obo/HAO_0001829 S3 http://purl.obolibrary.org/obo/HAO_0001831 S7 http://purl.obolibrary.org/obo/HAO_0002185 sculpture http://purl.obolibrary.org/obo/HAO_0000913 sternite http://purl.obolibrary.org/obo/HAO_0001654 sulcus http://purl.obolibrary.org/obo/HAO_0000978 T1 http://purl.obolibrary.org/obo/HAO_0000053 T2 http://purl.obolibrary.org/obo/HAO_0000056 T3 http://purl.obolibrary.org/obo/HAO_0000057 T7 http://purl.obolibrary.org/obo/HAO_0000061 tergite http://purl.obolibrary.org/obo/HAO_0001783 tibia http://purl.obolibrary.org/obo/HAO_0001017 tyloid http://purl.obolibrary.org/obo/HAO_0001199 vein http://purl.obolibrary.org/obo/HAO_0001095 vertex http://purl.obolibrary.org/obo/HAO_0001077 vertical epomial carina http://purl.obolibrary.org/obo/HAO_0000307 wing http://purl.obolibrary.org/obo/HAO_0001089

334 Acknowledgments Thanks to L. Musetti, J. Cora, and S. Hemly (OSUC) for critical assistance with specimen handling, software, and databasing; Agnièle Touret-Alby (MNHN) for imaging type material; Robert Zuparko (CAS) for the loan of type material. We thank the Florida Department of Agriculture and Consumer Services – Division of Plant Industry for their support on this contribution. This material is based upon work supported in part by the National Science Foundation of USA under grant No. DEB-0614764 to N.F. Johnson and A.D. Austin.

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336 Popovici O, Bin F, Masner L, Popovici M, Notton D (2011) Triteleia peyerimhoffi comb. n., a remarkably variable circum-Mediterranean scelionid (Hymenoptera, Platygastroidea). ZooKeys 140: 71–99. Risbec J (1950) Contribution à l'étude des Proctotrupidae (Serphiidae). Pages 511–639 in Risbec J. Travaux du Laboratoire d'Entomologie du Secteur Soudanis de Recherches Agronomiques. Gouvernement Général de l'Afrique Occidentale Française, Paris. 639 pp. Talamas EJ, Buffington ML (2015) Fossil Platygastroidea in the National Museum of Natural History, Smithsonian Institution. Journal of Hymenoptera Research 47: 1–52. Talamas EJ, Johnson NF, Buffington ML, Dong R (2016) Archaeoteleia Masner in the Cretaceous and a new species of Proteroscelio Brues (Hymenoptera, Platygastroidea). Journal of Hymenoptera Research 56: 241–261. Vilhelmsen L, Mikó I, Krogmann L (2010) Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insects: Hymenoptera). Zoological Journal of the Linnean Society 159: 22–194. Yoder MJ, Mikó I, Seltmann K, Bertone MA, Deans AR (2010) A gross anatomy ontology for Hymenoptera. PLosONE 5(12): e15991.

337 Chapter 4: A molecular reassessment of the phylogeny of the superfamily Platygastroidea (Hymenoptera)

Abstract Platygastroidea (Insecta: Hymenoptera) is one of the most diverse groups of parasitic wasps. All platygastroids are parasitoids of nine orders of insects or spiders. They appear to show a clear pattern of host group specificity. A robust phylogeny is essential to understanding the monophyly of and relationships among lower level groups and the pattern of shifts among host groups. Here, we conducted phylogenetic analyses of Platygastroidea based on four molecular markers (18S, 28S, COI, and wingless). Six major lineages are identified: 1) Archaeoteleia + Nixonia + Huddlestonium; 2) Platygastridae with Orwellium included; 3) Sparasion + Sceliomorpha; 4) the main scelionid clade; 5) Neuroscelio; and 6) Janzenella. Of the five previous recognized subfamilies, only Teleasinae and Telenominae s.s. are monophyletic. Relationships within the main scelionid clade are largely unresolved. Based on our results, the current classifications at familial, subfamilial, and tribal level within the superfamily are problematic. Without a robust phylogeny at hand, we reserve the consideration of a new family-level classification for the combined analyses of molecular and morphological data. Ancestral state reconstruction of host associations using maximum parsimony method supports that Orthoptera is the plesiomorphic host group for the superfamily. From this ancestral host group, a number of lineages of Platygastroidea have shifted to utilize Diptera, Hemiptera, Coleoptera, Araneae (spiders), Embioptera, Mantodea, Odonata, Lepidoptera and Neuroptera. The correlations between host shifts and radiations suggest causal relationships. To elucidate the underlying factors that determine the host range and host shifts among groups, a robust hypothesis of phylogenetic relationships is necessary. A combined analysis of additional sources of data (e.g., morphological and molecular characters) appears to hold the best prospects for future progress.

Introduction Over the past decade tremendous progress has been made in our understanding of the major outline of evolution and phylogeny in the insect order Hymenoptera (Sharkey 2007; Heraty et al. 2011; Sharkey et al. 2012; Klopfstein et al. 2013; Peters et al. 2017). The taxon is

338 made up of more than 150,000 described species as well as a large number of species yet to be formally described (Aguiar et al. 2013). The evolution of sociality within the order is the subject of intense scrutiny (Branstetter et al. 2017), but this behavior is restricted to a single clade within the order, the . Most Hymenoptera are parasitoids, a life history in which the immature stages feed upon, and eventually kill a single host . The most recent reconstruction of Hymenoptera phylogeny (Peters et al. 2017) proposed that the most diverse lineages of parasitoids – the Ceraphronoidea, , and Proctotrupomorpha – form a monophyletic group, dubbed the Parasitoida, that originated an estimated 236 mya. The Proctotrupomorpha itself is composed of three lineages, the Platygastroidea, , and a group composed of the superfamilies Proctotrupoidea, Diaprioidea, , and Chalcidoidea. The relationships among the three have not been firmly established. Our work is focused on the Platygastroidea, a diverse taxon with 6,041 species and 264 genera that currently are considered as valid. Most species are egg parasitoids of other insects and spiders, attacking hosts in the insect orders Odonata, Mantodea, Orthoptera, Embiidina, Hemiptera, Neuroptera, Lepidoptera, Coleoptera, and Diptera (Austin et al. 2005). One lineage, classically recognized as the Platygastridae, includes a relatively small number of species that are parasitoids of immature sternorrhynchous Hemiptera, and over 1000 species that are larval or egg-larval parasitoids of gall (Diptera: Cecidomyiidae). One unique feature of the superfamily is the presence of large multiporous basiconic sensilla on the ventral surface of the apical segments of the female antenna. These structures enable confident placement of fossil specimens in Platygastroidea. The oldest inclusions push the age of the superfamily back to at least 120 mya (Johnson et al. 2008). The traditional classification recognized two major taxa that largely reflect the differences in host relationships: the egg parasitoids in the family Scelionidae, and the gall fly parasitoids in the Platygastridae. To date there has been only a single phylogenetic analysis that has sought to sample the diversity across the entire superfamily. Murphy et al. (2007) analyzed the nucleotide sequences of three markers, 18S and 28S ribosomal DNA and mitochondrial cytochrome oxidase I, for >70 in-group species respresenting 55 genera of Platygastroidea. These data strongly support the hypothesis that the superfamily is a monophyletic group, and that within the superfamily the classical Platygastridae are also monophyletic. The latter group, however, arises from among genera that were traditionally grouped together as the Scelionidae,

339 thus rendering the scelionids as a paraphyletic taxon. On the basis of these results, Sharkey (2007) merged the two families into one, with the older name Platygastridae taking precedence. Within the superfamily 90% of the genera traditionally placed in the Scelionidae cluster together in a maximally-supported monophyletic group that Murphy et al. (2007) referred to as the “main scelionid clade.” The analysis supported the hypothesis of monophyly for only a few named higher taxa within the main clade: the tribe Scelionini and subfamilies Teleasinae and Telenominae. All other taxa above the rank of genus, at least those that are composed of and represented by more than one genus, appeared as para- or polyphyletic groups. This corroborated an earlier study of the spider egg parasitoids in the tribe Baeini (Carey et al. 2006). Subsequently, Taekul et al. (2014) also found that one subgroup of the Telenominae is more closely related to traditional members of the Scelioninae, concluding that even that well- defined subfamily was not monophyletic. Murphy et al. (2007) represented a significant, but preliminary step forward in understanding platygastroid phylogeny. The taxon sampling, 21% of known genera, did not include a number of taxa whose position within the old classification was unclear. The absence of a clear resolution of relationships within the main scelionid clade could also be the result of insufficient representation of lineages. Further, the analysis was based on sequences from rDNA and a mitochondrial gene: no nuclear protein-coding genes were included. The resulting collapse of much of the existing classification within Platygastroidea leaves little framework within which to interpret the evolution within the group, particularly habitat associations (e.g., the invasion of aquatic habitats) and host relationships. The goal of our work was to try to address the deficiencies in both taxon and gene sampling and thereby to test the conclusion of Murphy et al. (2007) and, hopefully, contribute to a clarification of relationships within the main clade. Materials and Methods Taxonomic sampling A total of 152 species belonging to 93 genera of Platygastroidea were included in the analyses, together with 4 outgroups comprising one species of each of , , , and Pelecinidae. The ingroup includes multiple representatives from all currently recognized platygastroid subfamilies and the majority of tribes. Molecular voucher specimens were freshly collected from around the world. After nondestructive DNA extraction

340 each voucher specimen was assigned with a unique barcode and deposited in the C. A. Triplehorn Insect Collection (OSUC). Voucher information and GenBank accession numbers for markers included in the analyses are available in Table 4.1. The metadata for the voucher specimens included in this study can be accessed in the Hymenoptera Online database (https://hol.osu.edu) using the identifiers (numbers prefixed with “OSUC” and USNMENT). A standard set of images were generated using JVC KY-F75U digital camera mounted in a Leica Z16 APOA microscope for most of the voucher specimens: dorsal habitus, lateral habitus, dorsal and lateral views of the head and mesosoma, and anterior view of head. The individual images are archived in Specimage (specimage.osu.edu) and are publically available.

DNA extraction, amplification, and sequencing Genomic DNA was extracted from ethanol-preserved specimens using the DNeasy Blood & Tissue Kit (Qiagen, Germantown, MD; cat. num. 69506) and following the protocol used by Taekul et al. (2014). Four gene regions were amplified using PCR: two nuclear ribosomal (18S and 28S D2-3), one mitochondrial protein encoding (COI), and one nuclear protein encoding locus (wingless). These genes were selected for their variability and resolving power at different levels in Hymenoptera (Heraty et al. 2011; Klopfstein et al. 2013). Primers for PCR amplification listed in Table 3.2. Two primers for amplification of an exon region of wingless, SceWgIF-1 and SceWgIR-1, were developed de novo for this study from existing Trissolcus basalis (Wollaston) genome sequences (Mao et al. 2012) using CLC Main Workbench. PCRs were carried out in 50 μL containing 25 μL ® GoTaq Green Master Mix, 2X (Promega, USA), 0.5 μL of 100 μM primers and 2.5 μL genomic DNA. Thermocycling conditions include initial denaturation at 94°C for 5 min, followed by 35 cycles of 1 min at 94 °C, 1 min at the primers’ annealing temperature, and 1.5 min of elongation at 72 °C, and ending with an additional extension of 72 °C for 3 min. PCR products were purified using the QIAquick PCR purification kit (Qiagen) protocol when necessary. Amplicons were directly sequenced in both directions with forward and reverse primers from Beckman Coulter Genomics (Danvers, MA). Chromatograms were assembled with Sequencher v4.0 (Gene Codes Corporation, Ann Arbor, MI). All the amplified sequences were deposited into GenBank (accession numbers begin with MF).

341 Sequence alignment and partitioning Ribosomal sequences of 18S and 28S were aligned with MAFFT version 7.309 (Katoh and Standley 2013) plugin in Geneious 11.0.3 using the E-INS-i algorithm with all parameters at their default values. This algorithm has been shown to be the preferred alignment algorithm for ribosomal sequences of Hymenoptera (Munro et al. 2011; Klopfstein et al. 2013). Protein-coding genes (COI and wingless) were aligned by codons using MUSCLE implemented in MEGA7 (Tamura et al. 2013). Exons were identified by alignment against known coding regions from GenBank. Introns are not objectively alignable (Klopfstein et al. 2013) and were removed from the final analyses.

Phylogenetic analyses Two phylogenetic estimation strategies were implemented to infer relationships: Maximum Likelihood (ML) and Bayesian inference (BI). ML analysis was performed using RAxML 8.2.10 (Stamatakis 2014), under the GTRCAT model carried out at the CIPRES website (https://www.phylo.org/). For this analysis, the concatenated alignment was partitioned by codon position of each protein-coding gene and each entire nuclear ribosomal gene. Rapid-bootstrap heuristic searches were calculated to estimate support levels from 1000 replicates. BI analysis was performed using MrBayes 3.2.6 (Ronquist et al. 2012) at the CIPRES website (https://www.phylo.org/). The concatenated data set was initially divided into 8 partitions: first, second, and third codon positions of COI and wingless, 18S, and 28S gene. The best-fitting nucleotide model for each partition was determined by the software PartitionFinderV1.1.1 (Lanfear et al. 2012) under the “greedy search” algorithm and the Bayesian information criterion (BIC) model metric (see Table 4.3). Two runs with four chains each were fun for a total of 150 million generations and sampled every 10 000 generations. The program Tracer 1.6 (Rambaut et al 2014) was used to check convergence and acceptable mixing. The majority rule consensus tree was generated after the initial 25% of saved trees were discarded as burn-in.

Phylogenetic informativeness tests To compare the phylogenetic performance of the four gene markers, we calculated their phylogenetic informativeness profiles using the online program PhyDesign

342 (http://phydesign.townsend.yale.edu/, López-Giráldez and Townsend 2011). This approach uses an ultrametric tree and an alignment to derive the evolutionary rate at which a character would be optimally phylogenetically informative for a given historical time on the phylogeny (Townsend 2007). The BI tree from the concatenated alignment was used the reference tree. Calculations were made with the concatenated alignment partitioned by gene (4 partitions, one for each gene), and by position (18S, 28S, three codon positions of COI and wingless).

Analysis of host association evolution Host association was chosen for Ancestral State Reconstruction (ASR). We were interested in the evolution of host associations among genera. Therefore, we used the ground-plan state of each genus for each terminal taxon. Each genus was scored based on the collecting information or information extracted from published literature and records stored in the Hymenoptera Online database (see Table 4.1). The genera of Gryon and have been reported to use several groups of host (e.g., Telenomus species are parasitoids of the eggs of Hemiptera, Lepidoptera, Diptera, and Neuroptera), but according to Taekul et al. (2014), the ground-plan hosts of Gryon and Telenomus are likely to be Hemiptera. The ancestral states of host association were reconstructed on the BI tree resulting from the analysis of concatenated data using both maximum parsimony (MP) and ML methods in MESQUITE v3.40 (Maddison & Maddison 2018).

Results Data properties In total, the nucleotide data matrix consisted of 3535 aligned sites for 152 ingroup taxa, of which 91 were represented by all loci; 17 lacked COI data; 53 lacked wingless data. Sequence summary statistics are given in Table 3.5 including information on total/average length and base frequencies. Analyses of overall phylogenetic informativeness for the four genes (Figure 4.1) ranked 28S as the most informative, followed by COI, wingless, and 18S, although the informativeness of 18S rose rapidly within a relative time of 0.4 but then declined steadily. From the by position analyses, the infromativeness of the 3rd codon of COI, the 1st and 2nd codon of wingless remained low throughout.

343

Phylogenetic relationships Both the BI and ML analyses of the concatenated data recovered similar well-supported clades. The phylogenetic trees resulting from the BI and ML analyses are shown in Figure 4.3 and Figure 4.4. The present analysis was not designed to study the included outgroups, but the results agree with previous studies (Murphy et al. 2007; Klopfstein et al. 2013; Peters et al. 2017). The monophyly of the Platygastroidea is highly supported (1.00 posterior probability (pp) and 73 bootstrap support (bs)). The relationships of the base of the tree are not well supported. Six groups form a polytomy. 1) Archaeoteleia, Nixonia, and Huddlestonium (0.86 pp, 58 bs): the support values for this clade are fairly low. Huddlestonium is represented by only two sequences (18S and 28S) and appears to render Nixonia paraphyletic. 2) Platygastridae is well supported as a monophyletic clade (1.00 pp, 89 bs) with Orwellium included, consistent with the results of Murphy et al. (2007). 3) Sparasion + Sceliomorpha (0.99 pp, 89 bs). 4) The main scelionid clade (1.00 pp, 70 bs), also consistent with Murphy et al. (2007). 5) Neuroscelio. 6) Janzenella. Within Platygastridae, the taxon sampling is not extensive. The Platygastrinae is not monophyletic: Allostemma is more closely related to Sceliotrachelinae. The monophyly of Sceliotrachelinae is only weakly supported in the BI analysis (0.80 pp). The ML analysis divides Platygastridae + Orwellium into the same two groups of genera as the BI analysis but the support values are quite low. Within the main scelionid clade both analyses support a clade composed of Probaryconus + Oethecoctonus at its base but the support values are low (0.75 pp, <50 bs). We can recognize some monophyletic groups, but relationships among them are largely unresolved. Telenominae sensu Taekul et al. (1.00 pp, 100 bs) and Teleasinae (1.00 pp, 100 bs) are maximally supported. Scelionini is well supported as monophyletic (1.00 pp) in the BI analysis. This tribe is weakly supported by the ML analysis (<50 bs), although a subset of the tribe (Pseudoheptascelio + Scelio + Synoditella) forms a monophyletic clade (99 bs). Other clades showing significant support include Baeini (sensu stricto, including Baeus, Ceratobaeus, Idris, and Odontacolus; 1.00 pp, 100 bs); Duta + Holoteleia + Masnerella + Spiniteleia (1.00 pp, 71 bs); the

344 Opisthacantha group with Ceratobaeus-type ovipositor (including Elgonia and Lapitha, 1.0 pp, 100 bs), and the Psix group sensu Taekul et al. (1.00 pp, 100 bs).

Host associations When host associations were mapped onto the BI tree from the concatenated data set, both MP and ML methods produced similar results (Figure 4.5; only the MP method tree is shown). Our results support that Orthoptera is the plesiomorphic host group for the superfamily as the basal groups such as Archaeoteleia, Nixonia and Sparasion are confined to orthopteran hosts. Many speciose genera within the main scelionid clade are also restricted to parasitize orthopteran eggs, for example, genera in the tribe Scelionini, Calliscelio, Paridris, and Macroteleia. Platygastridae use Coleoptera, Hemiptera and Diptera as hosts with Platygastrinae are restricted to attack dipterans (gall flies). Within the main scelionid clade, many genera of Scelioninae attack the eggs of Orthoptera but some groups have shifted to parasitize the eggs of other host groups, e.g., Baeini and Aradophagus parasitize the eggs of Araneae, Embidobia (Embidobiini) parasitize the eggs of Embioptera, Mantibarini and Axea parasitize the eggs of Mantodea, Calotelea and Thoronella parasitize the eggs of Odonata, Gryonini and the Psix group parasitize the eggs of Hemiptera. Teleasinae are restricted to parasitize the eggs of Coleoptera. The ancestral host associations in Telenominae are most likely Hemiptera, as suggested by Taekul et al. (2014).

Discussion

We reconstructed phylogenetic relationships among Platygastroidea based on sequences data of four genes (18S, 28S, COI, wingless) from 93 ingroup genera, which to date is the most extensive taxon sampling by far. Both the BI and ML analyses produced similar results (Figure 4.3, 4.4). The relationships among the basal groups within Platygastroidea are not well supported in our analyses, but six major groups are identified as described in the results. Our results support the monophyly of Platygastridae (with Orwellium included) as suggested by Murphy et al. (2007). Johnson et al. (2009) suggested that Orwellium was a sister group to Platygastrinae sensu lato (including both Platygastrinae and Sceliotrachelinae). The relationships between Orwellium

345 and the other groups within Platygastridae are not well supported in our analyses. Within Platygastridae, our results support the study of Murphy et al. (2007) that neither Platygastrinae nor Sceliotrachelinae is monophyletic, although Sceliotrachelinae is weakly supported as a monophyletic group in the BI analysis. Murphy et al. (2007) suggested that Archaeoteleia and Neuroscelio are basal to all other platygastroids. But the putative basal genera Janzenella and Nixonia were not included in their analyses. Our results show that Archaeoteleia, Nixonia, and Huddlestonium are weakly supported as a monophyletic group. Neuroscelio is not supported as a sister group to Archaeoteleia: rather it appears as a unique clade. The monophyly of Sparasion + Sceliomorpha was only weakly supported in the BI analysis in Murphy et al (2007), but is well supported in both analyses in this study. The position of Janzenella within the superfamily is unclear, as suggested by Masner and Johnson (2007) but appears to be a basal group. This genus is represented by only two sequences (18S and 28S) in this study. More sequences data may be helpful to resolve its relationships with other platygastroids. Our results support the monophyly of the main scelionid clade proposed by Murphy et al. (2007). The relationships within the main scelionid clade are not unclear. Scelioninae is not monophyletic, a result that is congruent with previous studies (Austin & Field 1997; Murphy et al. 2007). Our results support the monophyly of Teleasinae as suggested by Murphy et al. (2007). Murphy et al. (2007) recognized Telenominae as a monophyletic group, but this view was not supported by Taekul et al. (2014) because the Psix group was found to be more closely related to genera of Scelioninae. The remainder of Telenominae were supported as monophyletic as in the Taekul et al. (2014) analyses. The Psix group is also included in our analyses and our results are congruent with the conclusions of Taekul et al. (2014). When we examined the taxon sampling of these three studies, the monophyly of Telenominae concluded by Murphy et al. (2007) was due to the absence of the Psix group in their analyses. This implies that unrepresentative taxon sampling may affect the conclusions of phylogenetic relationships. As congruent with Murphy et al. (2007), most tribes within Scelioninae with multiple genera are not monophyletic. But we found some interesting relationships within Scelionini and Baeini, which are discussed below. Scelionini s.s. was only represented by two genera (Acanthoscelio and Scelio) in Murphy et al. (2007) study, and was weakly supported as monophyletic in their BI analysis. Six genera representing Scelionini are included in our analyses, and the tribe is recovered as monophyletic

346 in the BI tree. Dicroscelio was assigned in the tribe Scelionini by Masner (1976). Subsequently, Yoder et al. (2009) found that Dicroscelio sensu Masner was a new genus and described it as Axea Masner & Johnson, which was concluded as a misplacement within Scelionini. Yoder et al (2009) also found that Dicroscelio sensu Kieffer does not does not belong to Scelionini. Our results support the conclusions of Yoder et al. (2009): there is no evidence that neither Axea nor Dicroscelio are closely related to other genera in Scelionini. Baeini has been found to be non-monophyletic by Carey et al. (2006) and Murphy et al. (2007) because two genera of Baeini, Mirobaeoides and Neobaeus, are closely related to Embidobia. Our results support that Mirobaeoides is closely allied to Embidobia. However, our BI analysis weakly supports the affinity between Echthrodesis (Embidobiini) and the remainder of Baeini. Interestingly, all members of Baeini and Embidobiini are weakly supported as a monophyletic group in the BI analysis (0.71 pp), which was never recovered by previous studies (e.g., Carey et al. 2006, Murphy et al. 2007).

Implications for classification

Since the phylogeny published by Murphy et al. (2007), the classification in Platygastroidea at the family-level has been problematic. Without providing any new analysis of relationships, Sharkey (2007) synonymized Scelionidae with Platygastridae based on the paraphyly of Scelionidae concluded by Murphy et al. (2007), leaving a single family in Platygastroidea. This one family hypothesis is not useful. As pointed out by McKellar and Engel (2012), a classification pooling everything into Platygastridae s.l. would obscure much hierarchical as well as distinctive morphological and biological information. Subsequently, McKellar and Engel (2012) raised the putative basal lineages Nixoniini and Sparasionini to familial rank and proposed that Platygastroidea comprises four families: Nixoniidae, Sparasionidae, Scelionidae and Platygastridae. However, this four-familiy hypothesis is not without problems. First, the results of both Murphy et al. (2007) and this study suggest that Sparasionini is paraphyletic as Archaeoteleia is not closely related to other members of the tribe. If Archaeoteleia is not included in Sparasionidae, it is unplaced in the four-family scheme. Second, Nixoniidae contains only the putatively basal genus Nixonia, placed as such on the basis of its 14-merous antennae by Masner (1958). However, the two Nixonia species are grouped

347 together with Huddlestonium in our analyses. Huddlestonium has 13-segmented antennae, but is otherwise not similar to Nixonia. It is difficult to imagine morphological characters that could be used to recognize a taxon comprised of Huddlestonium (and the very similar genus Plaumannion) and Nixonia. Third, if Orwellium is included in Platygastridae, then the morphological definition of that family needs to be substantially revised. Finally, Neuroscelio and Janzenella are still unplaced in any family in the scheme of McKellar and Engel. The phylogenetic relationships presented in this study are not sufficient to propose a family-level classification. We reserve this consideration for the combined analyses of molecular and morphological data. At lower level of classification, these results are clearly insufficient to propose a comprehensive and stable subfamilial or tribal groupings.

Evolution of host associations in Platygastroidea Platygastroids use 9 orders of insects as well as spiders as hosts (Table 4.1), and a number of subfamilies and tribes are associated with a specific host group (Austin & Field 1997; Austin et al. 2005; Taekul et al. 2014). This pattern of host group specificity led Austin et al. (2005) to raise several interesting questions regarding the evolution of host associations. The absence of any phylogenetic scheme for higher-level taxa made it impossible to suggest answers. Host associations have been recorded for 52 of the 93 ingroup genera (56%) in this study. With the phylogeny presented in this study, we can begin to at least partly interpret the host relationships within the superfamily. Austin et al. (2005) suggested that Orthoptera is likely to be the plesiomorphic host group for the superfamily given that it is the most commonly utilized order across the Scelioninae and is the host group of the putatively basal genera Sparasion and Nixonia. Our current results support this hypothesis. The basal groups, including Archaeoteleia, Nixonia, and Sparasion are egg parasitoids of Orthoptera (Figure 4.5). In addition, a basal genus of the main scelionid clade, Oethecoctonus, also parasitizes orthopteran eggs. Although the relationships within the main scelionid clade are largely unresolved, orthopteran egg parasitism is commonly found in speciose genera of scelionine (Figure 4.5; Table 4.1). Subsequently, individual groups of platygastroid have undergone major changes from attacking one insect order to another or to spiders. Platygastridae have shifted to parasitize Diptera, Hemiptera, and Coleoptera. Platygastrinae were previously known only to parasitize gall

348 flies (Diptera). However, a recent study (Veenakumari et al. 2018) claimed that also parasitize psyllid species (Hemiptera). This record should be viewed with some caution because the Synopeas species was recorded from galls induced by a psyllid species and therefore the authors concluded that the psyllid was the host. But gall flies are equally likely the host, as was discussed by Austin (1984). Aradodophagus, all members of Baeini s. str., and Echthrodesis (Embidobiini) are parasitoids of spider eggs. As discussed above, Baeini and Embidobiini might be closely related. If this was the case, since eggs of both spiders and Embiidina are usually covered by silk, it suggests that these wasps may share a common evolutionary history of silk searching (Galloway & Austin 1984), and therefore use similar mechanisms to locate host eggs. Calotelea has been reported as parasitoids of Orthoptera and Odonata (Kononova & Fursov 2000), but is not close to Thoronella, which is only known to parasitize Odonata. Yoder et al. (2009) suspected that the host of Axea is Mantodea because one long series of specimens is labeled as “ex oothéque Phasmidae,” but walking sticks were not thought to produce Phasmatidae oothecae. A recent study, however, found that some Phasmatodea do produce oothecae (Goldberg et al. 2015). Therefore, the hosts for Axea could be either Mantodea or Phasmatodea. In either case, a close relationship between Axea and Mantibaria is not supported by our data. Thoron and Tiphodytes (Thoronini) are parasitoids of aquatic Hemiptera. Teleasinae are known only to parasitize the eggs of Coleoptera. Host shifts within Telenominae have been extensively discussed by Taekul et al. (2014). Briefly, the plesiomorphic hosts of these wasps appear to be terrestrial Hemiptera, from which multiple shifts have been made to attack to Lepidoptera, Diptera and Neuroptera (Taekul et al. 2014). In summary, over time, major groups within the superfamily have shifted from attacking Orthoptera to Diptera twice, at least twice to Coleoptera, spiders 2–3 times, Embiidina, Odonata 1–2 times, Mantodea 1–2 times, aquatic Hemiptera, terrestrial Hemiptera at least 3 times, Lepidoptera twice, and to Neuroptera once (Taekul et al. 2014). It is noteworthy that so far no single species is known to attack hosts in more than a single order. And at least half of these shifts are correlated with massive radiations of species. For example, Playgastrinae comprise more than 1800 species and are restricted to parasitize the eggs or of gall flies. Similarly, the shifts of Teleasninae to Coleoptera and of Gryonini + Telenominae to terrestrial bugs and then to Lepidoptera, Diptera and Neurotpera are correlated with radiations of each group into several hundreds or thousands of species.

349 The correlations between host shifts and radiations suggest a causal relationship. Abundant evidence has indicated that platygastroids rely on semiochemicals (e.g., sex and aggregation pheromones, cuticular hydrocarbons, female reproductive accessory gland secretions as well as plant-produced defensive chemicals) to find their hosts (Araraki et al. 1997; Bin et al. 1993; Colazza et al. 2004, 2007; Conti et al. 2004; Moraes et al. 2008; Peri et al. 2007; Salerno et al. 2009). Their chemosensory capacity may be a critical factor in shifting to a new group of hosts, and this may open the door to an explosion of new species exploiting this new resource. To critically explore this suggestion, a robust hypothesis of phylogenetic relationships is necessary. The results we present here are a modest step in that direction. The taxon sampling is fairly extensive in this study, and more taxon sampling seems unlikely to contribute to the resolution of relationships, at least those within the main scelionid clade. A combination of additional sources of data, such as morphological characters and the order of mitochondrial transfer RNA genes (Dowton et al. 2002), together with additional sequence data appear to hold the best prospects for future progress.

350 Table 4.1 Taxon sampling used in the molecular analysis listed in alphabetic order, including GenBank accession numbers. All GenBank accession numbers stating with MF and MH denote new sequences used for the first time in this study. The label data for all specimens have been geo‐ referenced and recorded in the Hymenoptera On‐ Line database. Details on the data associated with these specimens is publicly accessible at the following link, hol.osu.edu, and entering the specimen voucher ID in the form (e.g., OSUC 173847, note the space between the acronym and the number). Voucher Taxa 18S 28S COI wingless Host References IDs Ingroup Acantholapitha OSUC MF583199 MF583337 MF583475 MF583585 Unknown sp. 181548 Acanthoscelio OSUC MF583200 MF583338 MF583476 Unknown americanus 176021 Acanthoscelio OSUC MF583201 MF583339 MF583477 Unknown radiatus 181492 Acolomorpha OSUC MF583202 MF583340 MF583478 MF583586 Unknown sp. 165076 Allostemma sp. DQ888541 DQ888470 DQ888401 Unknown OSUC sp. MF583203 MF583341 MF583479 Diptera 203953 Vlug 1995 OSUC Amblyscelio sp. MF583204 MF583342 MF583480 MF583587 Unknown 181550 Masner & OSUC Amitus sp. MF583207 MF583345 MF583481 MF583588 Hemiptera Huggert 570566 1989 Anteris OSUC MF583208 MF583346 MF583483 MF583589 Unknown bilineata 339553 OSUC Apegus sp. MF583210 MF583348 MF583484 MF583590 Unknown 334009 Masner & Aphanomerus DQ888544 DQ888473 DQ888404 Hemiptera Huggert sp. 1989 OSUC Apteroscelio sp. MF583211 MF583349 MF583485 MF583591 Unknown 181537 Aradophagus OSUC Vetter et al. MF583214 MF583352 MF583487 MF583593 fasciatus 627833 Araneae 2012 Aradophagus OSUC Vetter et al. MF583212 MF583350 MF583592 sp. 534638 Araneae 2012 Archaeoteleia OSUC Austin & MF583215 MF583353 MF583488 MF583594 Orthoptera mellea 339570 Field 1997 Archaeoteleia Austin & DQ888505 JN623572 DQ888374 Orthoptera sp. Field 1997 OSUC Yoder et al. Axea sp. MF583216 MF583354 MF583489 MF583595 420745 Mantodea? 2009 OSUC Austin Baeus sp. MF583217 MF583355 MF583490 MF583596 627836 Araneae 1985 Muesebeck OSUC Baryconus sp. 1 MF583218 MF583356 MF583491 MF583597 Orthoptera 1979; Bin 420719 1976 Muesebeck OSUC Baryconus sp. 2 MF583220 MF583358 MF583492 Orthoptera 1979; Bin 176012 1976

351 Table 4.1 Continued Bracalba OSUC MF583221 MF583359 MF583493 MF583599 Unknown intermedia 230846 Chromoteleia OSUC MF583226 MF583364 MF583604 Unknown tricarinata 265262 Chromoteleia OSUC MF583224 MF583362 MF583495 MF583602 Unknown sp. 1 339616 Ceratobaeus OSUC Austin MF583228 MF583366 MF583496 MF583606 Araneae sp. 176054 1985 Cremastobaeus OSUC Austin & MF583229 MF583367 MF583497 MF583607 Orthoptera sp. 1 265205 Field 1997 Cremastobaeus OSUC Austin & MF583231 MF583369 MF583499 MF583609 Orthoptera sp. 2 534626 Field 1997 Calomerella OSUC MF583232 MF583370 MF583500 MF583610 Unknown scutellata 248143 Kononova OSUC Orthoptera; Calotelea sp. MF583234 MF583372 MF583502 MF583612 & Fursov 420731 Odonata 2000 Calliscelio OSUC Chen et al. JX303679 JX303701 JX303733 Orthoptera bisulcatus 176023 2017 Calliscelio OSUC Chen et al. MF583238 MF583376 MF583506 Orthoptera elegans 248095 2017 Calliscelio OSUC Chen et al. MF583237 MF583375 MF583505 Orthoptera longius 248070 2017 Calliscelio OSUC Chen et al. MF583235 MF583373 MF583503 MF583613 Orthoptera rubriclavus 534635 2017 OSUC Chen et al. Calliscelio sp. MF583236 MF583374 MF583504 MF583614 Orthoptera 420734 2017 OSUC Doddiella sp. MF583239 MF583377 MF583507 MF583615 Unknown 339548 OSUC Dichoteleas sp. MF583242 MF583380 MF583508 MF583618 Unknown 339610 OSUC Orthoptera; Dicroscelio sp. MF583243 MF583381 MF583509 HOL 266080 Hemiptera OSUC Masner Duta sp. 1 MF583244 MF583382 MF583510 Orthoptera 266066 1991 OSUC Masner Duta sp. 2 MF583245 MF583383 MF583511 MF583619 Orthoptera 181556 1991 OSUC Masner Duta sp. 3 MF583246 MF583384 MF583512 Orthoptera 188497 1991 Dyscritobaeus OSUC MF583247 MF583385 MF583513 Unknown sp. 176007 Echthrodesis OSUC van Noort MF583248 MF583386 MF583514 MF583620 Araneae lamorali 173844 et al. 2014 OSUC Embidobia sp. MF583249 MF583387 MF583621 Embioptera Dodd 1939 627843 Encyrtoscelio OSUC MF583250 MF583388 MF583622 Hemiptera Bin 1979 sp. 261873 OSUC Elgonia sp. MF583251 MF583389 MF583515 Unknown 181557 Eumicrosoma OSUC JX679552 JX683208 KC778481 Hemiptera Dicke 1937 sp. 173874 Masner & Fidiobia sp. DQ888545 DQ888474 DQ888405 Coleoptera Huggert 1989

352 Table 4.1 Continued Fusicornia OSUC MF583252 MF583390 MF583516 MF583623 Unknown spinosa 165072 OSUC Masner Gryon sp. 1 JX679535 JX683193 KC778430 MF583625 Hemiptera 266775 1983a OSUC Masner Gryon sp. 2 JX679536 JX683194 KC778432 MF583626 Hemiptera 266778 1983a OSUC Masner Gryon sp. 3 JX679537 JX683195 KC778429 MF583627 Hemiptera 266789 1983a OSUC Masner Gryon sp. 4 JX679538 JX683196 KC778433 Hemiptera 248210 1983a Gryonoides sp. OSUC MF583254 MF583392 MF583518 Unknown 1 266070 Gryonoides sp. OSUC MF583336 MF583474 MF583584 MF583708 Unknown 2 627839 Heptascelio OSUC Johnson et MF583257 MF583395 MF583519 MF583629 Orthoptera hamatus 192412 al. 2008 OSUC Johnson et Heptascelio sp. MF583256 MF583394 MF583628 Orthoptera 420732 al. 2008 Holoteleia OSUC MF583258 MF583396 Unknown bicolor 372800 Huddlestonium OSUC MF583259 MF583397 Unknown sp. 192430 OSUC Austin Idris sp. 1 MF583260 KF679327 KF679302 MF583630 Araneae 176058 1985 OSUC Austin Idris sp. 2 MF583261 MF583398 MF583520 MF583631 Araneae 420743 1985 OSUC Austin Idris sp. 3 MF583262 MF583399 MF583521 MF583632 Araneae 420751 1985 OSUC Austin Idris sp. 4 MF583263 MF583400 MF583522 MF583633 Araneae 534623 1985 Janzenella OSUC MF583255 MF583393 Unknown innupta 321942 Muesebeck sp. DQ888546 DQ888475 DQ888406 Diptera 1979 OSUC Lapitha sp. 1 MF583264 MF583401 MF583523 Unknown 188496 OSUC Lapitha sp. 2 MF583266 MF583403 MF583525 Unknown 248081 OSUC Masner Leptoteleia sp. MF583267 MF583404 MF583526 MF583635 Orthoptera 570554 1978 Macroteleia sp. OSUC Muesebeck MF583268 MF583405 MF583527 MF583636 Orthoptera 1 176053 1977 Macroteleia sp. OSUC Muesebeck MF583269 MF583406 MF583528 MF583637 Orthoptera 2 176026 1977 Macroteleia sp. OSUC Muesebeck MF583270 MF583407 MF583529 MF583638 Orthoptera 3 176011 1977 Mallateleia sp. DQ888530 DQ888456 DQ888394 Unknown OSUC Chopard Mantibaria sp. MF621555 MF621557 MF621560 Mantodea 248080 1923 OSUC Masnerella sp. MF583271 MF583408 MF583639 Unknown 570555 Masnerella OSUC MF583274 MF583411 MF583530 MF583642 Unknown cincta 181551

353 Table 4.1 Continued Mirobaeoides OSUC Austin MF583275 MF583412 MF583531 MF583643 Araneae sp. 339557 1985 OSUC Neoscelio sp. MF583276 MF583413 MF624485 MF583644 Unknown 570715 OSUC Neuroscelio sp. MF583278 MF583415 MF583532 Unknown 570714 Johnson & Nixonia OSUC MF583279 MF583416 MF583533 MF583646 Orthoptera Masner watshami 188493 2006 Johnson & OSUC Nixonia sp. MF583280 MF583417 MF583534 MF583647 Orthoptera Masner 248087 2006 Odontacolus OSUC Austin MF583281 MF583418 MF583535 MF583648 Araneae sp. 1 321852 1985 Odontacolus OSUC Austin MF583282 MF583419 MF583649 Araneae sp. 2 232369 1985 Oethecoctonus OSUC Masner MF583283 MF583420 MF583536 Orthoptera sp. 265217 1983b Opisthacantha OSUC MF583285 MF583422 MF583538 MF583651 Unknown sp. 534629 Oreiscelio OSUC MF583287 MF583424 MF583539 MF583653 Unknown naevus 207753 OSUC Orwellium sp. MF583288 MF583425 MF583540 Unknown 266096 OSUC Oxyteleia sp. JX679531 JX683189 KC778423 MF583654 Unknown 266063 OSUC Burks et al. Oxyscelio sp. MF583290 MF583427 MF583656 Orthoptera 372838 2013 OSUC Triapitsyn Parascelio sp. MF583291 MF583428 MF583541 MF583657 Hemiptera? 181575 et al. 2010 Paratelenomus OSUC Johnson JX679545 JX683206 KC778441 MF583660 Hemiptera saccharalis 270804 1985 Masner & OSUC Paridris gorn JX303684 JX303705 JX303718 MF583664 Orthoptera Huggert 405109 1968 Masner & OSUC Paridris mystax JX303668 JX303695 JX303714 MF583662 Orthoptera Huggert 254552 1968 Masner & OSUC Paridris nilaka JX303675 JX303694 JX303712 MF583661 Orthoptera Huggert 254613 1968 Masner & OSUC Paridris solaris JX303686 JX303706 JX303721 MF583663 Orthoptera Huggert 396849 1968 Masner & OSUC Paridris sp. MF583292 MF583429 MF583542 MF583665 Orthoptera Huggert 372837 1968 Phanuromyia OSUC Nesheim et JX679551 JX683210 KC778444 MF583666 Hemiptera sp. 270805 al. 2017 DQ888548 DQ888477 DQ888409 Unknown sp. sp. OSUC Muesebeck MF583332 MF583470 MF583704 Diptera 1 627831 1979

354 Table 4.1 Continued Platygaster sp. OSUC Muesebeck MF583333 MF583471 MF583583 MF583705 Diptera 2 627832 1979 Platygastoides DQ888549 DQ888478 DQ888410 Unknown sp. Platyscelidris OSUC MF583293 MF583430 MF583543 MF583667 Unknown fossorius 165081 Platyscelio OSUC Taekul et MF624484 MF583432 MF583545 Orthoptera pulchricornis 215797 al. 2010 Probaryconus OSUC MF583295 MF583433 MF583546 MF583668 Unknown rufipes 334146 Probaryconus OSUC MF583296 MF583434 MF583547 MF583669 Unknown sp. 570553 Johnson & Pseudoheptasce OSUC MF583297 MF583435 MF583548 Orthoptera Musetti lio muesebecki 176024 2011 Psilanteris sp. OSUC MF583298 MF583436 MF583549 MF583670 Unknown 1 266097 Psilanteris sp. OSUC MF583299 MF583437 MF583550 MF583671 Unknown 2 420730 Psilanteris sp. OSUC MF583300 MF583438 MF583551 MF583672 Unknown 3 176047 Johnson & OSUC Psix tunetanus JX679543 JX683200 KC778435 MF583673 Hemiptera Masner 266779 1985 Johnson & OSUC Psix sp. MF583301 MF583439 MF583552 MF583674 Hemiptera Masner 248211 1985 OSUC Romilius sp. MF583303 MF583441 MF583554 MF583676 Unknown 248084 OSUC Muesebeck Scelio striatus MF583305 MF583443 MF583556 MF583678 Orthoptera 203963 1972 Scelio OSUC Muesebeck MF583306 MF583444 MF583557 MF583679 Orthoptera zolotarevskyi 176009 1972 OSUC Muesebeck Scelio sp. MF583304 MF583442 MF583555 MF583677 Orthoptera 176075 1972 Sceliomorpha OSUC MF583307 MF583445 MF583558 MF583680 Unknown sp. 1 321940 Sceliomorpha OSUC MF583308 MF583446 Unknown sp. 2 176006 Sceliomorpha OSUC MF583309 MF583447 MF583559 MF583681 Unknown sp. 3 627835 OSUC Shreemana sp. MF583310 MF583448 MF583560 MF583682 Unknown 188492 USNME Sparasion Muesebeck NT0133 MH378417 MH378416 MH378415 Orthoptera cullaris 1979 5801 OSUC Muesebeck Sparasion sp. MF583311 MF583449 MF583561 Orthoptera 181549 1979 OSUC Spiniteleia sp. MF583312 MF583450 MF583562 MF583683 Unknown 372801 OSUC Masner Synoditella sp. MF583313 MF583451 MF583563 Orthoptera 165065 1976

355 Table 4.1 Continued Austin 1984; Diptera; Synopeas sp. DQ888551 DQ888480 DQ888411 Veenakum Hemiptera ari et al. 2018 OSUC Tanaodytes sp. MF583314 MF583452 Unknown 372788 OSUC Masner Thoron sp. MF583316 MF583454 MF583564 Hemiptera 248094 1972 OSUC Carlow Thoronella sp. MF583317 MF583455 MF583565 Odonata 176038 1992 Komeda et Teleas sp. DQ888560 DQ888488 Coleoptera al. 2015 Telenomus OSUC Johnson JX679572 JX683235 KC778462 MF583687 Hemiptera grenadensis 248194 1984 Telenomus OSUC Johnson JX679573 JX683236 KC778447 MF583684 Hemiptera nysivorus 266771 1984 Telenomus OSUC Johnson MF583318 MF583456 MF583566 MF583688 Hemiptera podisi 372849 1984 Telenomus sp. OSUC Johnson JX679582 JX683247 KC778473 MF583685 1 266780 Hemiptera 1984 Telenomus sp. OSUC Johnson JX679589 JX683256 KC778479 2 266781 Hemiptera 1984 Telenomus sp. OSUC Johnson JX679577 JX683241 KC778466 MF583686 3 266784 Hemiptera 1984 Telenomus sp. OSUC Johnson JX679595 JX683263 MF583567 MF583689 4 176014 Hemiptera 1984 Tiphodytes sp. OSUC Masner MF583319 MF583457 MF583568 Hemiptera 1 339736 1972 Tiphodytes sp. OSUC Masner MF583320 MF583458 MF583569 Hemiptera 2 176039 1972 OSUC Masner sp. 1 MF583321 MF583459 MF583570 MF583690 Diptera 372841 1983c OSUC Masner Trichacis sp. 2 MF583322 MF583460 MF583571 Diptera 372850 1983c Trichoteleia OSUC JX303673 JX303699 JX303728 Unknown hemlyae 334220 Trichoteleia OSUC JX303676 MF621558 MF583572 Unknown rugifrons 266092 Trichoteleia OSUC JX303671 JX303710 JX303725 Unknown tezitra 334221 Trichoteleia OSUC JX303670 JX303698 MF583573 MF583691 Unknown warren 254607 OSUC Muesebeck Trimorus sp. 1 MF583323 MF583461 MF583574 MF583692 Coleoptera 173837 1979 OSUC Muesebeck Trimorus sp. 2 MF583324 MF583462 MF583575 Coleoptera 176022 1979 OSUC Muesebeck Trimorus sp. 3 MF583334 MF583472 MF583706 Coleoptera 627837 1979 Trissolcus OSUC Talamas et JX679554 JX683212 KC778491 MF583694 Hemiptera hullensis 266782 al. 2015 Trissolcus OSUC Talamas et JX679555 JX683213 KC778486 MF583695 Hemiptera strabus 248187 al. 2015

356 Table 4.1 Continued Trissolcus OSUC Talamas et JX679553 JX683215 KC778487 MF583693 Hemiptera thyantae 266773 al. 2015 Trissolcus OSUC Johnson JX679556 JX683214 KC778485 MF583696 Hemiptera urichi 248199 1987 OSUC Talamas et Trissolcus sp. 1 JX679562 JX683225 KC778492 MF583697 Hemiptera 248205 al. 2015 OSUC Talamas et Trissolcus sp. 2 MF583325 MF583463 MF583576 MF583698 Hemiptera 372848 al. 2015 OSUC Trissoscelio sp. MF583326 MF583464 MF583577 Unknown 192433 OSUC Popovici et Triteleia sp. MF583253 MF583391 MF583517 MF583624 Orthoptera 214174 al. 2011 OSUC Zelandonota sp. MF583327 MF583465 MF583578 MF583699 Unknown 332135 Outgroup OSUC Megaspilus sp. MF621556 MF621559 MF621561 192624 Nasonia OSUC MF583329 MF583467 MF583580 MF583702 vitripennis 570561 Neodiprion OSUC MF583330 MF583468 MF583581 sertifer 570563 OSUC MF583331 MF583469 MF583582 MF583703 polyturator 553231

Table 4.2 Primers used to amplify four loci in this study.

Gene Primer name Primer sequences Reference 18S ai 5'-CCT GAG AAA CGG CTA CCA CAT C-3' Whiting et al. 1997 18S 18S-5R 5'-CTT GGC AAA TGC TTT CGC-3' Giribet et al. 1996 28S D2-3 D23F 5'-GAG AGT TCA AGA GTA CGT G-3' Whiting et al. 1997 28S D2-3 28Sb 5'-TCG GAA GGA ACC AGC TAC TA-3' Whiting et al. 1997 COI 1F 5'-GGA GGA TTT GGA AAT TGR YTW RTT CC-3' Simon et al. 1994 COI 1R 5'-ACT GTA AAT ATR TGA TGW GCT CA-3' Simon et al. 1994 COI Scel-f1 5'-GCA ATA ATT CGA ATA GAA TTA AGA GT-3' Gariepy et al. 2014 COI HCO-2198 5'-TAA ACT TCA GGG TGA CCA AAA AAT CA-3' Folmer et al. 1994 COI C1-J-1718 5'-GGA GGA TTT GGA AAT TGA TTA GTT CC-3' Simon et al. 2006 COI C1-N-2329 5'-ACT GTA AAT ATA TGA TGA GCT CA-3' Simon et al. 1994 wingless SceWgIF-1 5'-GTA AGT GTC ACG GGA TGT C-3' This study wingless SceWgIR-1 5'-TTG ACT TCA CAG CAC CAG T-3' This study wingless Wg_F 5'-GAR TGY AAR TGY CAY GGY ATG TCT GG-3' Brower & DeSalle 1998 wingless Wg_R 5'-ACT ICG CAR CAC CAR TGG AAT GTR CA-3' Brower & DeSalle 1998

357 Table 4.3 Substitution models identified by PartitionFinder for each partition.

Genetic maker Selected substitution model 18S TrN+I+G 28S SYM+I+G 1st codon 2nd codon 3rd codon position position position COI GTR+I+G TVM+I+G GTR+I+G wingless SYM+G TrNef+G SYM+I+G

Table 4.4 Summary statistics of genes used for phylogeny. Sequence length Base frequencies Genes # Taxa Shortest Longest Average %A %C %G %T 18S 152 533 957 673 24.8 21.9 28.1 25.1 28S 152 659 874 777 21.4 24.5 30.3 23.6 COI 135 192 645 577 33.2 15.2 12 39.6 wingless 99 375 489 467 23.2 26.4 29.6 20.9

358

Figure 4.1 Net phylogenetic informativeness profiles for the four genes used in this study.

359

Figure 4.2 Net phylogenetic informativeness profiles for the four genes and codon positions used in this study.

360

Figure 4.3 Relationships of Platygastroidea from Bayesian analysis of concatenated 18S, 28S, COI and wingless sequences with 8 partitions. Posterior probabilities above 0.50 indicated on branches.

361

Figure 4.4 Relationships of Platygastroidea from maximum likelihood analysis of concatenated 18S, 28S, COI and wingless sequences with 8 partitions. Boostrap values above 50 indicated on branches.

362

Figure 4.5 Host associations plotted on the BI tree (see tree Fig. 4.3) using maximum parsimony method in Mesquite.

363 Acknowledgments Thanks to Alejandro Valerio, Charuwat Taekul, and Elijah Talamas for collecting some of the molecular data and to Zachary Lahey and John V. Freudenstein for assistance in conducting phylogenetic analyses. This material is based upon work supported in part by the National Science Foundation of USA under grant No. DEB-0614764 to N.F. Johnson and A.D. Austin.

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368 Chapter 5: Identification and expression analysis of chemosensory receptor genes in the egg parasitoid Trissolcus basalis

Abstract Parasitic wasps are a diverse group of insects that have been widely used as biological control agents of insect pests in agriculture and forestry. These insects largely rely on their chemosensory abilities to locate food, mates, hosts, and other resources in a complex environment. Chemosensory receptors are thought to play a critical role in controlling the process of chemical perception in insects. Understanding the molecular evolution of chemosensory receptors in diverse groups of parasitic wasps can help us understand the chemosensory mechanisms that underlie host adaptation and speciation, which allows us to answer questions regarding species diversity. The parasitic wasp Trissolcus basalis is an effective biological control agent of the southern green stink bug, a pan-tropical agricultural pest, and is an ideal model for investigating the evolution of chemosensory receptors. We identified 34 gustatory receptors (Grs), 170 odorant receptors (Ors) and one odorant co-receptor (Orco), and 23 ionotropic receptors (Irs) in T. basalis through genomic and transcriptomic screening. Phylogenetic comparison with other hymenopteran species revealed differential Gr and Or expansions in T. basalis, including species-specific expansions of 21 Grs and 53 Ors. No species-specific expansions of Irs were found. Most of the species-specific expansions of Ors have 9 exons. Gender- and tissue-specific analyses showed that 5 Grs were only expressed in the antenna; 54 Ors were antenna-specific in both female and male, 66 Ors were antenna-specific only in female, and 4 Ors were antenna-specific only in male. No gender-specific Irs were found. The identification and expression profile of chemosensory receptor genes in T. basalis helps in understanding the link between the evolution of chemosensory receptors and speciation in parasitic wasps.

Introduction Parasitic wasps are important biological control agents of insect pests in agriculture and forestry (LaSalle & Gauld 1993; Asgari & Rivers 2011). In order to successfully suppress their hosts and reproduce, parasitic wasps rely on their chemosensory abilities to locate food, mates, hosts, and other resources in a complex environment. These abilities refer to the perception of

369 chemical cues such as those involved in smell (olfaction) and taste (gustation) (Suh et al. 2014). Chemical cues interact with a range of receptor proteins expressed in the sensory neurons (OSNs) of different types of chemosensory sensilla located on peripheral appendages such as the legs, wings, maxillary and labial palps, and especially the antennae (Suh et al. 2014; McKenzie et al. 2016). Among these proteins, the chemosensory receptor superfamily formed by the gustatory receptors (Grs), odorant receptors (Ors), and ionotropic receptors (Irs) are thought to be the most important (Zhou et al. 2015). However, despite the significant economic and ecological importance of parasitic wasps, only a few species have had their chemosensory receptor gene repertoires characterized (Robertson et al. 2010; Nishimura et al. 2012; Wang et al. 2015; Zhao et al. 2016). Further investigation on chemosensory receptors in diverse parasitic wasp groups is much needed. Much is known about the crucial roles that the major chemosensory receptor genes serve in insect chemosensory systems. Ors and Grs are seven-transmembrane domain receptors that belong to the same superfamily (Clyne et al. 1999, 2000). Classically, Ors are mainly expressed in the olfactory organs such as the antennae and maxillary palps where they detect volatile chemical signals including pheromones and general odorants (Touhara & Vosshall 2009). Functional olfactory receptors act primarily as ligand-gated ion channels and are composed of heteromeric complexes of two subunits: a highly conserved subunit now uniformly known as the olfactory receptor co-receptor (Orco) and a highly divergent subunit termed “tuning” Ors (Suh et al. 2014). The tuning Ors are responsible for odorant specificity, whereas Orco is required for proper function of tuning Ors (Stengl & Funk 2013). In contrast, most Grs are expressed in gustatory organs such as the proboscis, legs, and wings (Clyne et al. 2000; Scott et al. 2001; Dunipace et al. 2001), but some also in olfactory organs (Thorne & Amrein 2008). Grs respond to soluble tastants (e.g., sugars, bitter compounds, and salts) and some pheromones (Thorne et al. 2004; Vosshall & Stocker 2007; Freeman et al. 2014). However, one unusual group of Grs responds to volatile carbon dioxide (Jones et al. 2007). Irs belong to the most ancient family of chemoreceptors and are evolutionarily derived from ionotropic glutamate receptors (iGluRs) (Benton et al. 2009; Croset et al. 2010; Rytz et al. 2013). In Drosophila, Irs are expressed in coeloconic sensilla where Or/Orco complexes are usually absent (Scott et al. 2001). Two subfamilies are recognized within Irs: the conserved “antennal” Irs and the species-specific “divergent” IRs (Croset et al. 2010; Abuin et al. 2011). The antennal Irs are thought to act as

370 ligand-gated ion channels and respond to acid-based odorants or amines that are largely ignored by Ors (Yao et al. 2005; Abuin et al. 2011; Silbering et al. 2011). In contrast, the two divergent Irs, Ir8a and Ir25a, are thought to act as co-receptors, forming complexes with different sets of Irs to carry out odor-evoked neuronal responses (Croset et al. 2010; Abuin et al. 2011; Ai et al. 2013). The molecular features of chemosensory receptor genes can help to unravel the mechanisms of chemical perception from molecules to behavior in insects (Smadja and Butlin 2009). Therefore, an evolutionary comparison of these genes from different groups of insects may shed light on the chemosensory mechanisms of host adaptation and speciation in diverse insect taxa (Hallem et al. 2006). For example, chemoreceptors are one of the fastest evolving gene families in insects (Neafsey et al. 2015) and the expansion in one subfamily of odorant receptors is believed to play an important role in the evolution of sociality in honey bees (Robertson & Wanner 2006) and ants (Zhou et al. 2012; Engsontia et al. 2015; McKenzie et al. 2016). In addition, functional characterization of chemosensory genes may prove useful in the design of novel approaches to manipulate the behaviors of economically and medically important insects (Suh et al 2014). One key step prior to the functional characterization of chemosensory receptors is to determine the tissue-specific expression patterns. In insects, some chemosensory receptors have been found to be sex- and tissue-specific (Nishimura et al. 2012; Zhou et al. 2012). Therefore, it is critical to characterize the expression profiles of these genes in the major chemosensory organs of both sexes. The parasitic wasp Trissolcus basalis (Wollaston) is an ideal entry point for investigating the evolution of chemoreceptors. This species is an endoparasitoid that attacks the eggs of stink bugs and has been widely used as an effective biological control agent of the pan-tropical agricultural pest Nezara viridula (Linnaeus) (Hemiptera: Pentatomidae) in many countries (Loch & Walter 1999). This wasp uses multiple chemical cues to find mates (Loch & Walter 2002) and hosts (Mattiacci et al. 1993; Rosi et al. 2001; Colazza et al. 2009). In addition, the genus Trissolcus contains 193 described species, including both generalist and specialist species. Therefore, the identification of chemosensory receptor genes in T. basalis would help us understand the underlying mechanisms controlling chemical communication, behavior, host selection and acceptance, and host-wasp interactions so that we can design better biological

371 control strategies and develop monitoring tools to track parasitism rates in the field both for this species and related taxa. The goal of this research is to identify the chemosensory gene repertoires and characterize their molecular evolution in T. basalis via genome wide survey and transcriptomic analysis.

Materials and Methods Insect material, RNA extraction, and Illumina sequencing

The laboratory population of T. basalis used for this work originated from the southern United States. The colony was maintained on eggs of spined soldier bug, Podisus maculiventris (Say), which in turn were fed with mealworm, Tenebrio molitor Linnaeus, under laboratory conditions at 25 ±2 °C and 70±10% RH. The emerged adult wasps were collected within 24 h, immersed in RNA Later (Ambion, AM7020), and stored at -80 °C in Eppendorf tubes until further processing. The antennae of the wasps were dissected on dry ice. In total, 6 unit samples, 1000 antennae, 500 bodies without heads and 500 whole wasps of both sexes, were prepared for total RNA extraction using a Qiagen RNeasy Mini Kit (Qiagen, Hilden, Germany) following the manufacturer’s protocol. Total RNA was used to prepare a cDNA library using a TruSeq RNA library preparation kit v2 (Low-Throughput protocol; Illumina) following the manufacturer’s protocol. The quantity and quality of the libraries was estimated on an Agilent Bioanalyzer. Each library underwent paired-end sequencing using an Illumina HiSeq 2000 system at The Ohio State University’s Molecular and Cellular Imaging Center in Wooster, OH.

Transcriptome assembly and chemosensory receptor gene identification After removing short, low-quality sequences and adaptors, all reads from each sex and tissue were assembled separately using Trinity (Haas et al. 2013). The sequence homology search was performed using the BLASTX and BLASTN (Altschul et al. 1997) programs against the NCBI non-redundant protein and nucleotide sequence databases. Gene names were assigned to each unigene based on the best BLASTX hit with the highest score value. A TBLASTN search was performed using available chemosensory receptor gene sequences from Apis mellifera Linnaeus (Robertson & Wanner 2006), Nasonia vitripennis Walker (Robertson et al. 2010) and Microplitis demolitor Wilkinson (Zhou et al. 2012; Wang et al. 2015) as queries to

372 identify corresponding candidate unigenes in the transcriptomes of T. basalis (see Tables 5.1−5.3).

Annotation of T. basalis chemosensory receptor genes from the genome

To supplement the identification of chemosensory receptor genes from the transcriptomes, a genome-wide survey for these genes was performed. The draft genome of T. basalis was assembled using a combination of 454 and Illumina sequencing reads (Mao et al. 2012) and previous work in our lab has identified potential gene models for 36 Grs and 105 Ors (Talamas, unpublished). However, many of these genes were incomplete, indicating that the previous genome assembly needed to be improved. The genome of T. basalis was re-assembled by incorporating the previous draft genome with reads newly produced from Oxford Nanopore sequencing technologies. Chemosensory gene annotation followed the partially automated approach described in Zhao et al. (2012). Briefly, protein sequences of reported chemosensory genes collected from A. mellifera, M. demolitor, and N. vitripennis, as well as the newly discovered candidate unigens from the transcriptomes of T. basalis were used as queries for TBLASTN searches against the T. basalis genome database (http://macroteleia.biosci.ohio- state.edu/blast/blast.html) with an E-value cutoff of 1e-5. Putative chemosensory gene coding regions identified from the search results were subjected to gene model predictions in GeneWise (Birney et al. 2004) using the query amino acid sequence as a reference. All predictions inferred from GeneWise were manually checked for problem regions of models and pseudogenes in CLC Main Workbench and were refined according to the BLAST alignment. The BLAST and annotation process were iterated by adding newly predicted proteins from a previous run until no new genes were found. When the same gene identified from both the transcriptomes and the genome had different lengths, only the complete or the longer gene model was used in further analyses (see Tables 5.1−5.3).

Validation of chemosensory receptor gene models The final refined gene models were screened with InterProScan (Finn et al. 2017) to confirm the presence of characteristic domains of each protein family, and the conserved functions were determined by BLASTP in Geneious 11.0.3 against NCBI_NR database (see Tables 5.1−5.3).

373

Phylogenetic analysis For phylogenetic analyses of candidate genes of T. basalis, the data sets also included the receptor sequences from A. mellifera, M. demolitor, and N. vitripennis. The Gr data set comprised 34 amino acid sequences from T. basalis, together with A. mellifera (10), M. demolitor (79), and N. vitripennis (43). The Or data set comprised 171 amino acid sequences from T. basalis, together with A. mellifera (162), M. demolitor (203), and N. vitripennis (214). The Ir data set comprised 25 amino acid sequences from T. basalis, together with A. mellifera (11), M. demolitor (6), and N. vitripennis (11). The amino acid sequences of the candidate Grs, Ors, and Irs were aligned using the MAFFT (Katoh and Standley 2013) plugin in Geneious 11.0.3 with the FFT-NS-I algorithm, JTT200 scoring matrix and other default parameters. Phylogenetic analyses were performed using the Maximum likelihood (ML) method of PhyML3.0 (Guindon et al. 2010) with the best-fit model of evolution for each gene family was selected by SMS (Lefort et al. 2017) at the ATGC Montpellier Bioinformatics Platform (http://www.atgc-montpellier.fr). Rapid-bootstrap heuristic searches were calculated to estimate support levels from 100 replicates. For Ors, two rounds of phylogenetic analyses were conducted. First, all Ors of the four species were analyzed together. Second, all the T. basalis Ors were analyzed with selected representatives from the species-specific expansion clades of the other three hymenopteran species. The second round of analysis contains all the T. basalis Ors and 34, 27, and 39 Ors of A. mellifera, M. demolitor, and N. vitripennis, respectively.

Results Identification of chemosensory receptor genes All chemosensory receptor genes identified in T. basalis were named according to their orthology with those of A. mellifera, M. demolitor, and N. vitripennis, although this naming convention was inevitably disrupted due to gene loss or duplication in many of these taxa. The gene name, amino acid sequence length, status, scaffold number of the genome, number of exons, number of transmembrane domains, best Blast P hit, e-value, identity, and areas expressed are presented in Table 5.1–5.3 for Grs, Ors, and Irs, respectively. Amino acid sequences of all the putative chemosensory receptor genes are presented in Appendix A.

374 Gustatory receptors. Thirty-four candidate Grs were identified in T. basalis (Table 5.1). Of these, 16 are complete, 16 with the 5’ end partially lost, and two with the 3’ end partially lost. Eight of the 34 Grs were found in tandem arrays of 2–5 genes. The 16 complete Grs have 2–12 exons. The phylogenetic analysis of T. basalis Grs together with those from A. mellifera, M. demolitor, and N. vitripennis is shown in Figure 5.1. As in other Hymenoptera (Robertson & Wanner 2006; Robertson et al. 2010), we found no evidence of the highly conserved carbon dioxide receptors of Bombyx mori (Wanner & Robertson) and Diptera (De Bruyne et al 2003; Kent et al. 2008). Two Grs (TbGr1, TbGr2) were found in a clade with sugar receptors from A. mellifera, M. demolitor, and N. vitripennis. Of the remaining TbGrs, 21 T. basalis Grs (TbGr8, TbGr10, TbGr16–TbGr34) cluster in a species-specific subgroup, while the other TbGrs were grouped with the Grs of other hymenopteran species. The Gr gene expansions in T. basalis formed a lineage together with independent expansions of Grs in M. demolitor, and N. vitripennis. However, there is no evidence showing that this lineage is specific to the Parasitica because two A. mellifera Grs (AmGr4, AmGr5) are present in the expansions between N. vitripennis Grs and a subgroup of M. demolitor Grs. More than 50% of the Gr gene expansions in T. basalis are tandem arrays of 2–5 genes.

Odorant receptors. In total, 170 Ors and one copy of Orco were identified in T. basalis (Table 5.2). The TbOrco was recovered as complete and has a simple 1:1 orthology with known Orco in the NCBI database. Most of the Ors are complete except for 16 that are incomplete. Seventy-nine (46%) of the Ors are in tandem arrays of 2–5 genes. Of all the complete Ors, 129 (76%) were found complete in the genome assembly. These Ors have 2 to 12 exons, and 47 (36%) are 9- exon genes. A phylogenetic analysis of all T. basalis Ors together with representative Ors from A. mellifera, M. demolitor, and N. vitripennis is shown in Figure 5.2. A phylogenetic analysis of all Ors of the four species is shown in Figure 5.3. Like other hymenopteran species, the Or genes of T. basalis are more divergent than the Grs. With the exception of the Orco, most other relationships are complicated as to their orthology, and most involve at least 1: many, and often many : many relationships, indicating these clades of genes have undergone some degree of

375 duplication or loss. For example, there is an expansion of 18 TbOrs (TbOr23–TbOr40) with the single orthologue (AmOr35, MmOr29, NvOr5) from the other three hymenopteran species (Figure 5.2, 5.3). The expansions of TbOr41–TbOr48 and TbOr49–TbOr56 are grouped with different degrees of expansions of Ors in the other species. A species-specific expansion of 53 TbOrs (TbOr77–TbOr79, TbOr85–TbOr86, TbOr102–TbOr103, TbOr122–TbOr139, TbOr142–TbOr169) was also found in T. basalis. Most TbOrs in this expansion are 9-exon genes (Table 5.2).

Inotropic receptors Twenty-three candidate Irs and two N-methyl-D-aspartate (NMDA) receptors were identified in T. basalis (Table 5.3). Sixteen TbIrs are complete and 7 are partially lost at one or both ends. The 13 complete Irs identified from the genome have 3 to 19 exons. Only 2 genes, TbIr5 and TbIr9, are in tandem arrays. The phylogenetic analysis of Irs from the four hymenopteran species is shown in Figure 5.3. All T. basalis Irs are grouped with the Irs of the other hymenopteran species, including the two Ir co-receptors subfamilies: Ir8a (TbIr7), and Ir25a (TbIr5–TbIr6, TbIr8–TbIr10). No obvious species-specific expansions were found, although 4 Irs, TbIr1–TbIr4, are grouped with AmIr2 at the base of the tree.

Expression profile of chemosensory receptor genes in T. basalis Gustatory receptors. Of the 34 candidate Grs identified in T. basalis, 20 were retrieved from the transcriptomes (Table 5.1). The expression profile of Grs in T. basalis is shown in Figure 5.5. Five Grs are antenna-specific in both female and male, one Gr is body-specific to both female and male, and one Gr is male-specific in the whole wasp. The remaining Grs were found in the whole wasp of both female and male. No female-specific Grs were found.

Odorant receptors. Of the 170 Ors and the Orco identified in T. basalis, only TbOr54 was not retrieved from the transcriptomes (Table 5.2). The expression profile of Ors in T. basalis is shown in Figure 5.6. Fifty-four Ors are antenna-specific in both female and male, 66 Ors are antenna-specific

376 only in female, four Ors are antenna-specific only in male, two Ors are female-specific. The remaining Ors (including Orco) were found in the whole wasp of both female and male. Of the 66 female antenna-specific Ors, 31 (47%) are grouped in the T. basalis specific expansion of Ors, while none of the four male antenna-specific receptors is present in this Or expansion (Table 5.2 and Figure 5.2).

Ionotropic receptors All 23 Irs and 2 NMDA receptors identified in T. basalis were retrieved from the transcriptomes (Table 5.3). The expression profile of Irs in T. basalis is shown in Figure 5.7. Eight Irs (including one Ir co-receptor, TbIr5) are antenna-specific in both female and male, and only one Ir is body-specific. The remaining Irs were found in the whole wasp of both female and male. No gender-specific Irs were found.

Discussion

In this study, we identified 34 gustatory receptors (Grs), 170 odorant receptors (Ors) and one odorant co-receptor (Orco), and 23 inotropic receptors (Irs) in T. basalis both from the genome and transcriptomes. Considering the importance of T. basalis in biological control, the identified chemosensory receptor genes themselves provide valuable resources for future functional studies such as the identification of ligands for these receptors and the molecular mechanisms of chemoreception in governing host finding. More importantly, the comparative phylogenetic analyses of these chemosensory receptors among diverse parasitic wasp groups provide insights into their evolution and their roles in driving the linked processes of speciation and radiation in these important creatures. Only 20 of the 34 Grs identified from the T. basalis genome were retrieved from the transcriptomes. There could be several reasons for this finding. Grs are generally expressed at low levels (Biessmann et al. 2005; Thorne & Amrein 2008), and are mainly expressed in gustatory organs such as mouthparts, antennal dendrites, legs, and digestive tract. While the majority of Grs can be retrieved in the antennae, the expression of those in other parts of the body might be too low to be detected in the body transcriptomes. The expression of some Grs might be affected by the physiological condition of the parasitoids. The transcriptomic analyses were conducted on adult wasps in this study but some Grs might be expressed in other stages

377 such as larva and . It has been found that Or expression is amenable to modulation by scent conditioning in aphid endoparasitoid Aphidius gifuensis Ashmead (Hymenoptera: Braconidae) (Kang et al. 2017). In addition, some sugar taste receptors were differentially expressed in female parasitoids that reared on different host aphids. This suggests that taste conditioning might also modulate Gr expression. The wasps used in this study were freshly emerged adults, which had not been exposed to host eggs or plants. Some Grs might not be expressed because of the absence of taste stimulus. Some sugar taste Grs in A. gifuensis were found expressed highly in female than in male and this expression patterns might be related with different food types (Kang et al. 2017). Eight Grs in Chouioia cunea Yang (Hymenoptera: ) were found expressed much higher in female antennae than those in male (Zhao et al. 2016). These suggest that the differential expression patterns between females and males might be linked to the behaviors in locating mates and hosts. One Gr is found specific to males in T. basalis. This Gr might be related to finding female mates in males. We haven’t found any Grs are specific to T. basalis females. The fact that we did not find female-specific Grs implies: 1) Grs are not used by females to find hosts; or 2) higher expression level of Grs may mediate female-specific behaviors; or 3) these behaviors resulted from higher-level neural function and are not governed by peripheral sensory capacity. In order to identify the functions and ultimately their ligands of these Grs, more detailed studies of their expression pattern are needed. Ors are critical in sex pheromone and general odorants detections in insects (Touhara & Vosshall 2009). As in other hymenopterans, there is a large expansion of Ors than Grs in T. basalis. Almost all of the Ors in T. basalis can be retrieved in the transcriptomes and the majority of Ors is extensively expressed in the antennae. The conservative Orco is expressed in the whole wasp in both females and males, which is not surprising because Orco is required for the function of all Ors (Trible et al. 2017). Female wasps of T. basalis find fresh eggs to parasitize using chemical cues produced by the adult stage of their host (Conti & Colazza 2012, Fatouros et al. 2008). These include sex pheromones, aggregation pheromones, alarm pheromones, hydrocarbons deposited on plant surface, and secretions associated with oviposition. Compounds released by plants as a result of host-feeding or oviposition have also been shown to be biologically active (Conti & Colazza 2012). Male wasps are usually exposed to less complex chemical cues. Males usually emerge

378 one or two days before female and remain on their natal egg mass and mate with females immediately as they emerge (Austin et al. 2005). We identified 66 Ors are specific to female antennae, and only 4 Ors are specific to male antennae. This suggests that the female-specific Ors might be involved in the processes of locating and accepting hosts in females, while the male-specific Ors might be involved in the searching for female mates in males. Further studies are needed to test these hypotheses. Previous studies have found a large clade of Ors within the 9-exon Or family in social insects such as bees and ants and suggested that this expansion of Ors has been important in shaping social evolution in insects (Zhou et al. 2012; McKenzie et al. 2016, Karpe et al. 2017). However, according to our results, this hypothesis needs more extensive studies. We found an expansion of 53 Ors specific to T. basalis and most of these Ors are within the 9-exon Or family. Comparisons of gene intron evolution in the insect chemosensory receptors are thought to be informative (Robertson et al. 2010). More studies need to be carried out focusing on parasitic wasps. In this study, we identified 23 Irs in T. basalis. This number is greater than those in many studied parasitic wasps (for example, 11 in M. demolitor, 12 in N. vitripennis, 13 in Macrocentrus cingulum Brischke (Ahmed et al. 2016)), but is similar to A. gifuensis (Kang et al. 2017), which has 22. Irs are usually expressed in sensory neurons that respond to distinct odors but do not express either Ors or Grs (Benton et al. 2009). Compared with M. demolitor and N. vitripennis, T. basalis has relatively fewer Grs and Ors. The higher number of Irs found in T. basalis suggests that these Irs might supplement the capacity of this species to detect odors ignored by Grs or Ors. Chemical cues are known to be important in the process of host location and acceptance in parasitic wasps (Vinson 1976), but the details of the proximate mechanisms are not completely known for any species. Elucidation these mechanisms can help to better predict the host range of potential biological control agents and understand the role that host relationships play in the evolution and diversification of parasitic wasps. The chemosensory receptor genes identified in this study serve as the first step to investigate those mechanisms in the egg parasitoid T. basalis.

379 Table 5.1 Grs identified from the genome and transcriptomes in T. basalis. Numbers in square brackets indicate partial exons. FA, female antenna; FB, female body; MA, male antenna; MB, male body.

Gene ORF Exons Blast % Expressed Status Scaffold E-value name (aa) (no.) P hit Identity in XP_019887704 PREDICTED: gustatory FA, FB, TbGr1 457 Complete 1062 10 receptor for sugar taste 64f isoform X8 0.00E+00 62 MA, MB [Ooceraea biroi] XP_014232824 gustatory receptor for FA, FB, TbGr2 461 Complete 1062 9 sugar taste 64f-like [Trichogramma 1.06E-172 59 MA, MB pretiosum] XP_012276873 putative gustatory FA, FB, TbGr3 411 Complete 814 7 1.58E-59 31 receptor 28b, partial [Orussus abietinus] MA, MB XP_015597026 PREDICTED: putative FA, FB, TbGr4 435 Complete 770 12 gustatory receptor 28a isoform X2 6.91E-32 27 MA, MB [Cephus cictus] XP_014219005 gustatory receptor for FA, FB, TbGr5 497 Complete 235 8 sugar taste 43a-like [Copidosoma 2.00E-144 44 MA, MB floridanum] XP_006614814 PREDICTED: putative FA, FB, TbGr6 399 Complete 1974 7 8.84E-81 39 gustatory receptor 28b-like [Apis dorsata] MA, MB NP_001177438 gustatory receptor12 TbGr7 309 5' Lost 754 [3] 6.66E-13 25 [Nasonia vitripennis] XP_011875522 PREDICTED: putative TbGr8 336 5' Lost 6058 [3] gustatory receptor 28a [Vollenhovia 4.44E-04 24

emeryi] NP_001177438 gustatory receptor 12 FA, FB, TbGr9 335 Complete 770 2 1.39E-15 23 [Nasonia vitripennis] MA, MB NP_001177440 gustatory receptor 14 TbGr10 318 5' Lost 754 [3] 4.14E-11 23 [Nasonia vitripennis] XP_014611281 PREDICTED: putative FA, FB, TbGr11 381 5' Lost 551 [4] gustatory receptor 23a, isoform B 2.54E-96 42 MA, MB [Polistescanadensis] XP_012229639 PREDICTED: putative TbGr12 377 5' Lost 551 [5] gustatory receptor 28a [Linepithema 3.67E-67 35 FA, MA humile] XP_012229639 PREDICTED: putative TbGr13 408 5' Lost 551 [2] gustatory receptor 28a [Linepithema 1.54E-105 43

humile] XP_015109675 PREDICTED: putative FA, FB, TbGr14 409 Complete 551 3 gustatory receptor 28b [Diachasma 2.95E-121 48 MA, MB alloeum] XP_015109675 PREDICTED: putative FA, FB, TbGr15 402 Complete 551 3 gustatory receptor 28b [Diachasma 1.11E-121 48 MA, MB alloeum] XP_012229082 PREDICTED: putative TbGr16 389 Complete 340 4 gustatory receptor 28a isoform X1 1.24E-33 27 FA, MA [Linepithema humile] XP_012229082 PREDICTED: putative TbGr17 397 Complete 887 4 gustatory receptor 28a isoform X1 9.48E-13 26 FB, MB [Linepithema humile] XP_012273861 putative gustatory FA, FB, TbGr18 388 Complete 962 4 5.87E-27 27 receptor 28a [Orussus abietinus] MA, MB

380 Table 5.1 Continued KZC10989 Putative gustatory receptor TbGr19 416 Complete 328 5 4.90E-34 27 FA, MA 28b [Dufourea novaeangliae] OAD61494 Putative gustatory receptor TbGr20 325 3' Lost 1280 [2] 5.18E-06 29 28b [Eufriesea mexicana] KYN09746 Putative gustatory receptor FA, FB, TbGr21 400 Complete 1280 4 2.13E-24 28 28b [Trachymyrmex cornetzi] MA, MB XP_019885475 PREDICTED: putative TbGr22 397 Complete 4137 5 gustatory receptor 28b isoform X1 1.92E-25 27 FA, MA [Camponotus floridanus] XP_020292579 putative gustatory TbGr23 328 5' Lost 505 [4] receptor 28b isoform X1 [Pseudomyrmex 4.94E-11 24

gracilis] KZC10988 Putative gustatory receptor TbGr24 389 5' Lost 505 [4] 3.14E-29 28 MA, MB 28b [Dufourea novaeangliae] XP_019888560 PREDICTED: putative TbGr25 382 Complete 2882 2 1.67E-36 31 FA, MA gustatory receptor 28a [Ooceraea biroi] KYM88343 Putative gustatory receptor TbGr26 325 5' Lost 4284 [1] 7.83E-04 24 28b [Atta colombica] XP_012536589 PREDICTED: putative FA, FB, TbGr27 340 3' Lost 426 [1] gustatory receptor 28a [Monomorium 1.37E-06 28 MA, MB pharaonis] XP_012536589 PREDICTED: putative TbGr28 313 5' Lost 3239 [1] gustatory receptor 28a [Monomorium 4.40E-07 24

pharaonis] XP_011640850 PREDICTED: putative TbGr29 305 5' Lost 4284 [1] gustatory receptor 28b isoform X3 8.55E-12 23

[Pogonomyrmex barbatus] XP_011640848 PREDICTED: putative TbGr30 309 5' Lost 3239 [1] gustatory receptor 28a isoform X1 1.28E-12 23

[Pogonomyrmex barbatus] XP_015119835 PREDICTED: gustatory TbGr31 348 5' Lost 3239 [2] receptor for bitter taste 66a-like 1.17E-07 24

[Diachasma alloeum] XP_011165419 PREDICTED: putative TbGr32 328 5' Lost 4284 [1] gustatory receptor 28a [Solenopsis 3.48E-10 25

invicta] XP_015119835 PREDICTED: gustatory TbGr33 316 5' Lost 3239 [1] receptor for bitter taste 66a-like 2.79E-09 27

[Diachasma alloeum] KYN16346 Putative gustatory receptor TbGr34 348 5' Lost 3239 [1] 2.46E-08 25 28b [Trachymyrmex cornetzi]

381 Table 5.2 Ors identified from the genome and transcriptomes in T. basalis. Numbers in square brackets indicate partial exons. FA, female antenna; FB, female body; MA, male antenna; MB, male body.

Gene ORF Exons Blast % Expressed Status Scaffold E-value name (aa) (no.) P hit Identity in NP_001310774 odorant receptor FA, FB, TbOrco 480 Complete 1076 8 0.00E+00 78 coreceptor [Cephus cinctus] MA, MB XP_014226814 LOW QUALITY TbOr1 414 Complete NA NA PROTEIN: odorant receptor 22c- 6.53E-36 26 FA like [Trichogramma pretiosum] NP_001177643 odorant receptor FA, FB, TbOr2 398 Complete 49 5 2.66E-120 45 288 [Nasonia vitripennis] MA, MB NP_001177643 odorant receptor TbOr3 413 Complete 285 5 4.09E-101 42 FA, MA 288 [Nasonia vitripennis] XP_012224802 PREDICTED: FA, FB, TbOr4 396 Complete 49 [4] odorant receptor 67c-like 7.60E-109 42 MA, MB [Linepithema humile] XP_016837003 PREDICTED: TbOr5 388 Complete 2133 6 odorant receptor 289 isoform X1 1.25E-139 51 FA, MA [Nasonia vitripennis] XP_011347234 PREDICTED: FA, FB, TbOr6 336 5' Lost 36 4 odorant receptor 33b-like isoform 3.51E-19 26 MA, MB X2 [Ooceraea biroi] XP_016836982 PREDICTED: FA, FB, TbOr7 401 Complete 2325 4 odorant receptor 41 isoform X1 1.45E-55 28 MA, MB [Nasonia vitripennis] XP_011347241 PREDICTED: TbOr8 365 Complete 1193 4 odorant receptor 13a [Ooceraea 9.53E-27 24 FA biroi] NP_001177434 odorant receptor FA, FB, TbOr9 406 Complete 385 5 4.62E-24 25 8 [Nasonia vitripennis] MA, MB XP_020295550 odorant receptor TbOr10 397 Complete 994 5 4.19E-35 26 FA, MA 4-like [Pseudomyrmex gracilis] XP_015603609 PREDICTED: FA, FB, TbOr11 437 Complete 415 4 odorant receptor 4-like isoform 3.27E-30 25 MA, MB X1 [Cephus cinctus] XP_006565953 PREDICTED: FA, FB, TbOr12 400 Complete 1746 NA odorant receptor 4-like isoform 2.60E-23 26 MA, MB X2 [Apis mellifera] XP_012214672 PREDICTED: TbOr13 400 Complete 455 4 odorant receptor 4-like 2.05E-35 26 FA, MA [Linepithema humile] XP_015603609 PREDICTED: FA, FB, TbOr14 414 Complete 455 5 odorant receptor 4-like isoform 4.19E-33 26 MA, MB X1 [Cephus cinctus] XP_015603609 PREDICTED: FA, FB, TbOr15 432 Complete 455 5 odorant receptor 4-like isoform 1.17E-42 28 MA, MB X1 [Cephus cinctus] XP_017880563 PREDICTED: TbOr16 316 Complete 642 2 odorant receptor 4-like [Ceratina 1.26E-18 24 FA, MA calcarata] XP_003700932 PREDICTED: FA, FB, TbOr17 402 Complete 6246 5 odorant receptor Or2-like 1.81E-34 27 MA, MB [Megachile rotundata]

382 Table 5.2 Continued XP_003700932 PREDICTED: TbOr18 406 Complete 1391 2 odorant receptor Or2-like 1.93E-35 26 MA [Megachile rotundata] XP_003394006 odorant receptor FA, FB, TbOr19 406 Complete 2763 4 3.63E-35 26 4-like [Bombus terrestris] MA, MB XP_003394006 odorant receptor TbOr20 410 Complete NA NA 1.67E-31 27 FA 4-like [Bombus terrestris] XP_020279260 odorant receptor TbOr21 405 Complete 178 5 1.98E-36 25 FA 4-like [Pseudomyrmex gracilis] XP_011880220 PREDICTED: TbOr22 397 Complete 642 6 odorant receptor 49b-like 3.49E-175 59 FA, MA [Vollenhovia emeryi] XP_015112584 PREDICTED: FA, FB, TbOr23 391 Complete 4739 5 odorant receptor 67c-like 4.44E-61 29 MA, MB [Diachasma alloeum] XP_015112584 PREDICTED: TbOr24 430 Complete 711 4 odorant receptor 67c-like 1.37E-81 37 MA [Diachasma alloeum] XP_015112584 PREDICTED: FA, FB, TbOr25 415 Complete 42 3 odorant receptor 67c-like 3.36E-112 42 MA, MB [Diachasma alloeum] XP_015112584 PREDICTED: TbOr26 425 Complete 711 4 odorant receptor 67c-like 2.20E-88 35 FA, MA [Diachasma alloeum] XP_015112584 PREDICTED: FA, FB, TbOr27 422 Complete 711 5 odorant receptor 67c-like 1.12E-88 36 MA, MB [Diachasma alloeum] XP_020279256 odorant receptor FA, FB, TbOr28 372 Complete 192 4 3.90E-44 25 82a-like [Pseudomyrmex gracilis] MA, MB XP_018310069 PREDICTED: TbOr29 379 Complete 192 5 odorant receptor 22c-like 1.09E-38 28 FA, MA [Trachymyrmex zeteki] XP_014604750 PREDICTED: TbOr30 379 Complete 192 6 odorant receptor 45b-like 1.17E-50 30 FA, MA [Polistes canadensis] XP_015112584 PREDICTED: TbOr31 323 Complete 192 5 odorant receptor 67c-like 2.33E-34 24 FA, MA [Diachasma alloeum] XP_015112584 PREDICTED: TbOr32 395 Complete NA NA odorant receptor 67c-like 7.69E-73 33 FA, MA [Diachasma alloeum] XP_015112584 PREDICTED: TbOr33 400 Complete 5432 4 odorant receptor 67c-like 2.71E-68 33 FA, MA [Diachasma alloeum] XP_015112584 PREDICTED: TbOr34 328 Complete 264 4 odorant receptor 67c-like 2.49E-50 31 FA, MA [Diachasma alloeum] XP_003393917 odorant receptor TbOr35 394 Complete 524 NA 1.51E-52 28 FA 67a-like [Bombus terrestris] XP_015112584 PREDICTED: TbOr36 394 Complete 3578 7 odorant receptor 67c-like 6.98E-53 28 FA, MA [Diachasma alloeum]

383 Table 5.2 Continued XP_015431139 PREDICTED: TbOr37 448 Complete 3578 4 odorant receptor 33b-like 5.80E-49 29 FA, MA [Dufourea novaeangliae] XP_017795948 PREDICTED: FA, FB, TbOr38 398 Complete 3578 4 odorant receptor 67a-like 2.37E-63 32 MA, MB [Habropoda laboriosa] XP_017795970 PREDICTED: TbOr39 398 Complete 3578 5 odorant receptor 4-like 3.52E-61 33 FA, MA [Habropoda laboriosa] XP_015431139 PREDICTED: TbOr40 392 Complete 3578 7 odorant receptor 33b-like 1.42E-64 33 FA, MA [Dufourea novaeangliae] NP_001177492 odorant receptor FA, FB, TbOr41 376 Complete 1745 6 7.86E-66 33 45 [Nasonia vitripennis] MA, MB XP_014299871 PREDICTED: TbOr42 371 Complete 1745 6 odorant receptor 49b-like 1.82E-54 31 FA, MA [Microplitis demolitor] NP_001177492 odorant receptor FA, FB, TbOr43 388 Complete 2291 4 1.10E-71 34 45 [Nasonia vitripennis] MA, MB NP_001177492 odorant receptor FA, FB, TbOr44 379 Complete 1745 6 9.50E-71 34 45 [Nasonia vitripennis] MA, MB NP_001177492 odorant receptor FA, FB, TbOr45 365 Complete 1745 5 9.94E-70 33 45 [Nasonia vitripennis] MA, MB NP_001177492 odorant receptor TbOr46 369 Complete 601 6 6.32E-51 27 FA, MA 45 [Nasonia vitripennis] NP_001177492 odorant receptor TbOr47 368 Complete 346 6 6.08E-51 28 FA, MA 45 [Nasonia vitripennis] NP_001177492 odorant receptor TbOr48 372 Complete 346 6 7.92E-45 26 FA, MA 45 [Nasonia vitripennis] AKO90005 odorant receptor 41 TbOr49 371 Complete 3357 6 3.28E-37 30 FA, MA [Microplitis mediator] KYN03550 Putative odorant TbOr50 376 Complete 1012 6 receptor 49a [Cyphomyrmex 5.83E-32 27 FA, MA costatus] XP_015182349 PREDICTED: TbOr51 373 Complete 1800 6 odorant receptor 43a-like isoform 5.14E-50 29 FA, MA X1 [Polistes dominula] NP_001177491 odorant receptor TbOr52 375 Complete 2207 6 1.13E-50 28 FA, MA 44 [Nasonia vitripennis] XP_014601080 PREDICTED: FA, FB, TbOr53 372 Complete 5720 2 odorant receptor 30a-like 7.43E-49 30 MA, MB [Polistes canadensis] NP_001177491 odorant receptor TbOr54 233 3', 5' Lost 5720 [2] 2.98E-26 27 44 [Nasonia vitripennis] AKO90005 odorant receptor 41 TbOr55 199 3' Lost 2819 [2] 8.50E-29 30 FA, MA [Microplitis mediator] NP_001177491 odorant receptor TbOr56 209 3', 5' Lost 5720 NA 1.54E-24 27 FA, MA 44 [Nasonia vitripennis] XP_015177210 PREDICTED: TbOr57 378 Complete 4145 [2] odorant receptor 13a-like 5.16E-101 41 FA, MA [Polistes dominula] XP_006567985 PREDICTED: FA, FB, TbOr58 377 Complete 881 [5] odorant receptor 13a-like isoform 4.93E-87 36 MA, MB X2 [Apis mellifera]

384 Table 5.2 Continued ALR87041 odorant receptor 141 FA, FB, TbOr59 401 Complete 3094 4 1.52E-77 35 [Apis cerana cerana] MA, MB XP_012529718 PREDICTED: TbOr60 410 Complete 372 5 odorant receptor 13a-like 1.74E-46 29 MA [Monomorium pharaonis] XP_012529718 PREDICTED: TbOr61 401 5' Lost 3871 [6] odorant receptor 13a-like 3.18E-58 31 MA [Monomorium pharaonis] AKO89978 odorant receptor 14 FA, FB, TbOr62 428 Complete 666 6 1.54E-63 31 [Microplitis mediator] MA, MB AKO89978 odorant receptor 14 FA, FB, TbOr63 412 Complete 642 6 2.66E-78 34 [Microplitis mediator] MA, MB XP_024222684 odorant receptor FA, FB, TbOr64 401 Complete 890 6 2.10E-37 27 13a-like [Bombus impatiens] MA, MB XP_017796295 PREDICTED: TbOr65 401 Complete 890 [6] odorant receptor 30a-like 1.80E-29 30 FA, MA [Habropoda laboriosa] XP_012344900 PREDICTED: FA, FB, TbOr66 356 Complete 890 6 odorant receptor 13a-like [Apis 2.39E-29 28 MA, MB florea] XP_011691922 PREDICTED: FA, FB, TbOr67 406 Complete 1025 6 odorant receptor Or2-like 7.22E-90 38 MA, MB [Wasmannia auropunctata] XP_014608934 PREDICTED: TbOr68 402 Complete 859 6 odorant receptor Or2-like 1.79E-59 31 FA, MA [Polistes canadensis] XP_015183883 PREDICTED: FA, FB, TbOr69 404 Complete 1305 5 odorant receptor 30a-like 8.53E-53 31 MA, MB [Polistes dominula] XP_014608934 PREDICTED: TbOr70 397 Complete 3514 6 odorant receptor Or2-like 6.43E-58 28 FA, MA [Polistes canadensis] XP_015183884 PREDICTED: TbOr71 404 Complete 506 6 odorant receptor 49b-like 2.75E-82 38 FA, MA [Polistes dominula] XP_014295110 PREDICTED: TbOr72 386 Complete 1968 5 odorant receptor 83a-like 1.98E-18 24 FA [Microplitis demolitor] NP_001164395 odorant receptor FA, FB, TbOr73 395 Complete 71 4 1.61E-97 39 82 [Nasonia vitripennis] MA, MB NP_001164419 odorant receptor FA, FB, TbOr74 392 Complete 1150 5 9.35E-46 28 249 [Nasonia vitripennis] MA, MB NP_001164419 odorant receptor FA, FB, TbOr75 391 Complete 1150 5 2.79E-51 30 249 [Nasonia vitripennis] MA, MB NP_001177594 odorant receptor FA, FB, TbOr76 381 Complete 1150 6 2.06E-54 28 245 [Nasonia vitripennis] MA, MB XP_012538364 PREDICTED: TbOr77 435 Complete 984 8 odorant receptor 4-like isoform 1.44E-43 26 FA X1 [Monomorium pharaonis] XP_020721192 odorant receptor TbOr78 398 Complete 146 9 3.25E-21 24 FA, MA 9a [Bombus terrestris] XP_015108891 PREDICTED: TbOr79 408 Complete 146 9 odorant receptor 10a-like 3.17E-30 24 FA, MA [Diachasma alloeum]

385 Table 5.2 Continued XP_019882173 PREDICTED: TbOr80 394 Complete 400 9 odorant receptor 13a 4.63E-05 21 FA [Camponotus floridanus] XP_012248015 uncharacterized TbOr81 407 Complete 1132 [4] protein LOC105681630 isoform 1.42E-03 26 FA X4 [Bombus impatiens] NP_001310780 odorant receptor FA, FB, TbOr82 396 Complete 1058 9 5.60E-09 23 Or2-like [Cephus cinctus] MA, MB XP_020277684 odorant receptor FA, FB, TbOr83 385 Complete 3034 NA 8.53E-23 23 13a-like [Pseudomyrmex gracilis] MA, MB AQN78460 olfactory receptor 58 TbOr84 374 Complete 818 8 3.92E-18 22 FA [Meteorus pulchricornis] XP_012529131 PREDICTED: FA, FB, TbOr85 411 Complete 945 9 odorant receptor 24a-like 1.81E-21 25 MA, MB [Monomorium pharaonis] XP_012138711 PREDICTED: TbOr86 410 Complete 254 9 odorant receptor Or2-like isoform 1.26E-15 22 FA X3 [Megachile rotundata] XP_011703994 PREDICTED: TbOr87 272 5' Lost 5933 NA odorant receptor 13a-like isoform 1.43E-09 26 FA X5 [Wasmannia auropunctata] XP_012282483 odorant receptor TbOr88 397 Complete 2368 9 4-like isoform X3 [Orussus 1.31E-07 23 FA abietinus] XP_016770140 PREDICTED: TbOr89 397 Complete 859 9 odorant receptor 98 isoform X1 8.60E-07 29 FA, MA [Apis mellifera] XP_003700936 PREDICTED: FA, FB, TbOr90 419 Complete 3151 8 odorant receptor 13a-like 1.10E-06 24 MA, MB [Megachile rotundata] XP_019565304 PREDICTED: TbOr91 404 Complete 4299 NA odorant receptor 83a-like [Aedes 9.33E-07 22 FA, MA albopictus] XP_012143140 PREDICTED: TbOr92 407 Complete 3454 [6] odorant receptor 49a-like 1.89E-06 25 FA [Megachile rotundata] KOX67782 Odorant receptor Or2 TbOr93 406 Complete 1622 [8] 4.19E-08 25 FA, FB [Melipona quadrifasciata] XP_012537251 PREDICTED: TbOr94 403 Complete 1622 9 odorant receptor 4-like 1.20E-07 22 FA [Monomorium pharaonis] XP_011883592 PREDICTED: TbOr95 403 Complete 6330 11 odorant receptor 13a-like 2.48E-10 29 FA [Vollenhovia emeryi] XP_019884411 PREDICTED: FA, FB, TbOr96 391 Complete 1793 10 odorant receptor 49b isoform X2 1.03E-10 30 MA, MB [Camponotus floridanus] XP_016845697 PREDICTED: TbOr97 414 Complete 111 9 odorant receptor 132 isoform X1 3.65E-12 22 FA [Nasonia vitripennis] XP_012145209 PREDICTED: TbOr98 382 Complete 2456 10 odorant receptor 43a-like 4.69E-06 29 FA, MA [Megachile rotundata]

386 Table 5.2 Continued XP_012055093 PREDICTED: TbOr99 380 Complete 5570 8 odorant receptor 85c-like [Atta 1.71E-04 22 FA cephalotes] XP_017095092 PREDICTED: TbOr100 372 Complete 6726 9 odorant receptor 2a [Drosophila 1.47E-08 22 FA bipectinata] NP_001177516 odorant receptor TbOr101 353 Complete 6726 9 3.98E-07 25 FA, MA 93 [Nasonia vitripennis] AII01107 odorant receptor TbOr102 411 Complete 3318 9 2.60E-14 28 FA [Dendrolimus kikuchii] XP_017795547 PREDICTED: TbOr103 415 Complete 3318 9 odorant receptor 9a-like 5.24E-22 24 FA, MA [Habropoda laboriosa] XP_011882831 PREDICTED: odorant receptor 46a, isoform A- TbOr104 400 Complete 3892 9 4.26E-12 24 FA like isoform X1 [Vollenhovia emeryi] XP_011330820 PREDICTED: TbOr105 263 Complete 6821 [2] odorant receptor Or1-like isoform 3.40E-09 27 FA X1 [Ooceraea biroi] AJO62236 olfactory receptor TbOr106 402 Complete 3892 9 2.08E-11 22 FA OR17 [Tenebrio molitor] AJO62224 olfactory receptor TbOr107 398 Complete 4404 [4] 5.47E-11 20 FA OR5 [Tenebrio molitor] XP_012255495 odorant receptor TbOr108 383 Complete 1128 10 85b-like isoform X1 [Athalia 1.75E-09 20 FA rosae] XP_020279052 odorant receptor TbOr109 308 5' Lost 792 [7] 2.87E-09 30 FA, MA Or2-like [Pseudomyrmex gracilis] XP_011500883 PREDICTED: TbOr110 403 Complete 4555 10 odorant receptor 56a [Ceratosolen 1.36E-04 35 FA solmsi marchali] NP_001310772 odorant receptor TbOr111 297 3', 5' Lost 2629 [7] 4.18E-09 21 FA, MA 13a-like [Cephus cinctus] NP_001177575 odorant receptor TbOr112 397 Complete 2307 [6] 2.97E-09 23 FA, MA 203 [Nasonia vitripennis] NP_001177513 odorant receptor TbOr113 394 Complete 994 10 1.28E-06 26 FA, MA 87 [Nasonia vitripennis] XP_018406443 PREDICTED: TbOr114 398 Complete 3404 [7] odorant receptor 4-like 2.91E-08 21 FA [Cyphomyrmex costatus] XP_012537251 PREDICTED: TbOr115 396 Complete 2004 9 odorant receptor 4-like 2.24E-12 26 FA [Monomorium pharaonis] XP_021203888 odorant receptor TbOr116 399 Complete 190 9 1.91E-06 22 FA 4-like [Bombyx mori] XP_011869950 PREDICTED: TbOr117 398 Complete 4125 8 odorant receptor Or1-like isoform 2.20E-07 26 FA X1 [Vollenhovia emeryi] XP_011146335 PREDICTED: TbOr118 398 Complete 3103 9 odorant receptor 46a-like isoform 2.68E-07 25 FA X2 [Harpegnathos saltator]

387 Table 5.2 Continued XP_017778204 PREDICTED: TbOr119 398 Complete 4628 10 putative odorant receptor 92a 2.15E-06 28 FA [Nicrophorus vespilloide] XP_012537251 PREDICTED: TbOr120 398 Complete 1930 9 odorant receptor 4-like 6.64E-08 26 FA [Monomorium pharaonis] XP_012531579 PREDICTED: TbOr121 369 Complete 1110 8 odorant receptor 33a-like 4.88E-04 28 FA [Monomorium pharaonis] XP_020721192 odorant receptor TbOr122 399 3' Lost 1652 [9] 9.57E-20 25 FA 9a [Bombus terrestris] XP_012169049 odorant receptor TbOr123 401 Complete 4302 9 9a isoform X1 [Bombus 2.71E-10 22 FA, MA terrestris] XP_012169049 odorant receptor TbOr124 421 5' Lost NA NA 9a isoform X1 [Bombus 4.36E-09 21 FA, MA terrestris] XP_011301634 PREDICTED: TbOr125 385 Complete 235 9 odorant receptor 13a-like [Fopius 1.49E-09 21 FA, MA arisanus] XP_014298635 PREDICTED: TbOr126 391 Complete 273 [5] odorant receptor 67c-like 4.26E-17 25 FA [Microplitis demolitor] XP_008549292 PREDICTED: TbOr127 396 Complete 273 10 odorant receptor 22c-like 1.19E-11 20 FA [Microplitis demolitor] XP_015171022 PREDICTED: TbOr128 426 Complete 2907 11 odorant receptor 9a-like [Polistes 3.47E-17 26 FA dominula] XP_017758779 PREDICTED: TbOr129 412 Complete 273 9 odorant receptor 4-like [Eufriesea 4.23E-07 25 FA, MA mexicana] NP_001177555 odorant receptor TbOr130 427 Complete 273 11 2.54E-09 22 FA 167 [Nasonia vitripennis] XP_015108891 PREDICTED: TbOr131 416 Complete 2682 12 odorant receptor 10a-like 2.37E-11 23 FA [Diachasma alloeum] XP_011334606 PREDICTED: TbOr132 414 Complete 7042 9 odorant receptor 22c isoform X1 1.66E-08 21 FA [Ooceraea biroi] XP_011309690 PREDICTED: TbOr133 421 Complete 5966 9 odorant receptor 13a-like isoform 2.30E-07 20 FA X1 [Fopius arisanus] NP_001177541 odorant receptor TbOr134 391 Complete 190 9 6.93E-15 22 FA 137 [Nasonia vitripennis] XP_011871787 PREDICTED: FA, FB, TbOr135 397 Complete 51 [7] odorant receptor 4-like 1.49E-15 26 MA, MB [Vollenhovia emeryi] XP_019699736 PREDICTED: FA, FB, TbOr136 400 Complete 51 [6] odorant receptor 22c-like isoform 2.80E-18 24 MA, MB X4 [Harpegnathos saltator] XP_016839048 PREDICTED: TbOr137 390 Complete 3318 9 odorant receptor 43a [Nasonia 4.83E-16 23 FA vitripennis]

388 Table 5.2 Continued NP_001177548 odorant receptor FA, FB, TbOr138 408 Complete 599 9 1.31E-23 25 147 [Nasonia vitripennis] MA, MB AQN78423 olfactory receptor 21 FA, FB, TbOr139 386 Complete 599 9 9.76E-11 26 [Meteorus pulchricornis] MA, MB XP_012537251 PREDICTED: TbOr140 410 Complete 3318 9 odorant receptor 4-like 7.83E-18 24 FA [Monomorium pharaonis] XP_008202544 PREDICTED: TbOr141 408 Complete 3318 9 odorant receptor 140 isoform X1 2.12E-23 24 FA [Nasonia vitripennis] NP_001177702 odorant receptor TbOr142 404 Complete 1137 9 4.21E-12 22 FA, MA 156 [Nasonia vitripennis] XP_014297094 PREDICTED: TbOr143 411 Complete 1137 9 odorant receptor 22a-like 7.29E-18 22 FA [Microplitis demolitor] XP_011309690 PREDICTED: TbOr144 391 Complete 4822 [4] odorant receptor 13a-like isoform 1.55E-19 24 FA X1 [Fopius arisanus] XP_011878580 PREDICTED: TbOr145 412 Complete 1670 9 odorant receptor 4-like isoform 1.34E-20 23 FA X3 [Vollenhovia emeryi] XP_020285667 odorant receptor TbOr146 248 5' Lost NA NA 2.24E-11 25 FA, MA 13a-like [Pseudomyrmex gracilis] XP_019882173 PREDICTED: TbOr147 401 Complete 1312 [7] odorant receptor 13a 4.63E-05 21 FA [Camponotus floridanus] XP_011878580 PREDICTED: TbOr148 410 5' Lost 2695 [9] odorant receptor 4-like isoform 6.78E-12 21 FA X3 [Vollenhovia emeryi] AGG17940 olfactory receptor 7 TbOr149 431 Complete 2069 9 4.15E-14 22 FA [Microplitis mediator] XP_019699550 PREDICTED: TbOr150 409 Complete 2069 9 odorant receptor 4 isoform X4 2.95E-16 24 FA [Harpegnathos saltator] XP_020280604 odorant receptor TbOr151 409 Complete 399 9 1.46E-09 22 FA 13a-like [Pseudomyrmex gracilis] XP_024228077 uncharacterized TbOr152 406 5' Lost 399 [9] protein LOC100741411 [Bombus 2.09E-10 23 FA impatiens] XP_012543307 PREDICTED: TbOr153 410 Complete 2875 9 odorant receptor 63a-like 1.74E-06 21 FA [Monomorium pharaonis] NP_001177702 odorant receptor TbOr154 306 Complete 1633 6 2.45E-08 27 FA, FB 156 [Nasonia vitripennis] XP_011865384 PREDICTED: TbOr155 417 5' Lost 187 [10] odorant receptor 22c-like isoform 3.55E-22 22 FA X1 [Vollenhovia emeryi] NP_001177543 odorant receptor TbOr156 349 Complete 1745 7 2.49E-16 27 FA, MA 140 [Nasonia vitripennis] XP_006562271 PREDICTED: TbOr157 421 Complete 2712 9 odorant receptor Or1-like [Apis 3.66E-13 22 FA, MA mellifera] NP_001177545 odorant receptor TbOr158 412 Complete 358 9 1.81E-16 23 FA 143 [Nasonia vitripennis]

389 Table 5.2 Continued XP_019884288 PREDICTED: putative odorant receptor 85d TbOr159 353 3' Lost 28 [9] 8.47E-21 25 FA isoform X1 [Camponotus floridanus] XP_012224104 PREDICTED: TbOr160 411 Complete 28 9 odorant receptor 45b-like isoform 1.03E-18 25 FA, MA X1 [Linepithema humile] XP_012529131 PREDICTED: TbOr161 379 Complete 28 9 odorant receptor 24a-like 1.10E-12 24 FA, MA [Monomorium pharaonis] NP_001177545 odorant receptor TbOr162 398 Complete 3369 9 1.80E-05 22 FA 143 [Nasonia vitripennis] XP_016837025 PREDICTED: TbOr163 376 Complete 1975 7 odorant receptor 141 isoform X1 7.74E-16 21 FA [Nasonia vitripennis] XP_011138659 PREDICTED: TbOr164 409 Complete 1975 9 odorant receptor 9a-like isoform 1.88E-10 20 FA X1 [Harpegnathos saltator] XP_012531307 PREDICTED: TbOr165 409 Complete NA NA odorant receptor 13a-like isoform 1.51E-10 21 FA, MA X1 [Monomorium pharaonis] XP_011259871 PREDICTED: TbOr166 400 Complete 3369 [6] odorant receptor 63a-like 1.05E-12 20 FA [Camponotus floridanus] XP_012531307 PREDICTED: TbOr167 415 Complete 1975 9 odorant receptor 13a-like isoform 1.42E-07 19 FA X1 [Monomorium pharaonis] KYN03370 Odorant receptor 22b TbOr168 310 3' Lost 2584 [9] 2.00E-06 25 FA [Cyphomyrmex costatus] XP_019696169 PREDICTED: TbOr169 405 Complete 2584 9 odorant receptor 24a-like 9.23E-08 23 FA [Harpegnathos saltator] XP_015181320 PREDICTED: FA, FB, TbOr170 401 Complete 352 9 odorant receptor 13a-like 1.66E-81 35 MA, MB [Polistes dominula]

390 Table 5.3 Irs identified from the genome and transcriptomes in T. basalis. FA, female antenna; FB, female body; MA, male antenna; MB, male body.

Gene ORF Exons Blast % Expressed Status Scaffold E-value name (aa) (no.) P hit Identity in PBC30154 Glutamate [NMDA] receptor subunit [Apis cerana FA, FB, TbNmdar1 1038 Complete 717 6 cerana] 0 70 MA, MB XP_012537361 PREDICTED: glutamate receptor ionotropic, NMDA 2B isoform X4 TbNmdar2 947 Complete 408 13 [Monomorium pharaonis] 0 83 FB, MB XP_015590333 PREDICTED: glutamate receptor ionotropic, kainate 2-like isoform X1 [Cephus FA, FB, TbIr1 1019 Complete 1484 19 cinctus] 0 85 MA, MB XP_012147937 PREDICTED: glutamate receptor ionotropic, kainate 2-like isoform X2 [Megachile FA, FB, TbIr2 900 Complete 1026 14 rotundata] 0 55 MA, MB XP_015522026 PREDICTED: glutamate receptor ionotropic, TbIr3 690 Complete 3224 13 kainate 2-like [Neodiprion lecontei] 0.00E+00 81 Fb, MB XP_011152744 PREDICTED: glutamate receptor ionotropic, FA, FB, TbIr4 915 Complete 1710 17 kainate 2 [Harpegnathos saltator] 0.00E+00 54 MA, MB TbIr5 XP_023316789 ionotropic receptor (Ir25a.1) 477 5' Lost 3113 NA 25a [Trichogramma pretiosum] 1.37E-100 36 FA, MA TbIr6 XP_012287489 ionotropic receptor FA, FB, (Ir25a.2) 683 5' Lost 615 NA 25a [Orussus abietinus] 5.37E-53 25 MA, MB XP_011685531 PREDICTED: TbIr7 glutamate receptor ionotropic, FA, FB, (Ir8a) 909 Complete 9 17 kainate 2 [Wasmannia auropunctata] 0 63 MA, MB XP_012522547 PREDICTED: TbIr8 glutamate receptor ionotropic, delta-1 FA, FB, (Ir25a.3) 365 3', 5' Lost 6036 NA [Monomoriumpha raonis] 9.23E-116 45 MA, MB XP_015585435 PREDICTED: TbIr9 glutamate receptor ionotropic, FA, FB, (Ir25a.4) 452 Complete 3113 3 kainate 4 [Cephus cinctus] 6.82E-173 55 MA, MB XP_015585435 PREDICTED: TbIr10 glutamate receptor ionotropic, FA, FB, (Ir25a.5) 928 5' Lost 456 NA kainate 4 [Cephus cinctus] 0 58 MA, MB XP_015598828 PREDICTED: glutamate receptor ionotropic, kainate 3 isoform X1 [Cephus TbIr11 857 Complete 659 13 cinctus] 0 36 FA, MA TbIr12 XP_012265004 glutamate receptor FA, FB, (Ir76b) 522 Complete 181 8 ionotropic, delta-1 [Athalia rosae] 1.43E-133 43 MA, MB XP_018052627 PREDICTED: ionotropic receptor 25a-like isoform FA, FB, TbIr13 640 Complete 1456 NA X1 [Atta colombica] 0 46 MA, MB XP_012539815 PREDICTED: glutamate receptor 1-like TbIr14 617 Complete 2513 8 [Monomorium pharaonis] 2.51E-55 27 FA, MA

391 Table 5.3 Continued XP_014295944 PREDICTED: glutamate receptor ionotropic, kainate 5, partial [Microplitis TbIr15 299 3', 5' Lost 3151 NA demolitor] 8.25E-21 24 FA, MA KZC09836 Glutamate receptor delta- FA, FB, TbIr16 668 Complete 61 9 2 subunit [Dufourea novaeangliae] 0.00E+00 49 MA, MB XP_020718982 glutamate receptor ionotropic, kainate 2 [Bombus TbIr17 640 Complete 3680 9 terrestris] 2.07E-119 35 FA, MA XP_015191547 PREDICTED: glutamate receptor 1-like [Polistes FA, FB, TbIr18 403 5' Lost 2124 NA dominula] 2.72E-93 42 MA, MB XP_020718982 glutamate receptor ionotropic, kainate 2 [Bombus FA, FB, TbIr19 671 Complete 6922 NA terrestris] 6.61E-116 33 MA, MB XP_014214903 glutamate receptor ionotropic, kainate 1-like TbIr20 509 5' Lost NA NA [Copidosoma floridanum] 2.35E-108 38 FA, MA AQN78503 ionotropic receptor 64a TbIr21 621 Complete 990 10 [Meteorus pulchricornis] 1.16E-175 44 FA, MA ARN17852 ionotropic receptor 6 FA, FB, TbIr22 629 Complete 226 10 [Cephus cinctus] 0 49 MA, MB ARN17852 ionotropic receptor 6 TbIr23 619 Complete 7380 NA [Cephus cinctus] 0 47 FA, MA

392

Figure 5.1 Maximum likelihood phylogenetic tree of gustatory receptors (Grs) from T. basalis and other hymenopteran speices. Boostrap values above 50 indicated on branches. Am, A. mellifera, black; Mm, M. demolitor, orange; Nv, N. vitripennis, blue; Tb, T. basalis, red.

393

Figure 5.2 Maximum likelihood phylogenetic tree of all odorant receptors (Ors) from T. basalis and representatives of Ors from other hymenopteran speices. Boostrap values above 50 indicated on branches. Am, A. mellifera, black; Mm, M. demolitor, orange; Nv, N. vitripennis, blue; Tb, T. basalis, red.

394

Figure 5.3 Maximum likelihood phylogenetic tree of all odorant receptors (Ors) from T. basalis and other hymenopteran speices. Boostrap values above 50 indicated on branches. Am, A. mellifera, black; Mm, M. demolitor, orange; Nv, N. vitripennis, blue; Tb, T. basalis, red.

395

Figure 5.4 Maximum likelihood phylogenetic tree of inotropic receptors (Irs) from T. basalis and other hymenopteran speices. Boostrap values indicated on branches. Am, A. mellifera, black; Mm, M. demolitor, orange; Nv, N. vitripennis, blue; Tb, T. basalis, red.

396 MA+MB, 1

FB+MB, 1

FA+MA, 5

FA+FB+MA+MB, 13

Figure 5.5 Tissue- and gender-specific expression profile of Grs in T. basalis. FA: female antenna; FB: female body; MA: male antenna; MB: male body.

397 MA, 4

FA+FB+MA+MB, 44

FA, 66

FA+FB, 2

FA+MA, 54

Figure 5.6 Tissue- and gender-specific expression profile of Ors in T. basalis. FA: female antenna; FB: female body; MA: male antenna; MB: male body.

398 FA+MA, 8

FA+FB+MA+MB, 14

FB+MB, 1

Figure 5.7 Tissue- and gender-specific expression profile of Irs in T. basalis. FA: female antenna; FB: female body; MA: male antenna; MB: male body.

399 Acknowledgments Thanks to George Keeney for assistance in maintaining insect material, to Elijah J. Talamas for identifying some of the gene models from the genome, and to Zachary Lahey for assistance in conducting phylogenetic analyses. This material is based upon work supported in part by the National Science Foundation of USA under grant No. DEB-0614764 to N.F. Johnson and A.D. Austin.

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416 Appendix A: amino acid sequences of TbGrs, TbOrs, and TbIrs in FASTA format

>TbGr1 MRKLVCKNGKKSQVMDEPLNRVFNTNMDIVSSEVIRPVISNRENYVTNTDSLHLALRPII IMAQCFGLFPVSGINSPDTSHLSFTWKSFKIVYNCIAALSVLVLTVYSIHRLTTTNFTSNK TSNVVFYATTGLTSILFIKLAKKWPNFATSWESIERELSTRHPRNNSRISLALRFKIISSVV MTLALLEHGLSILSGIRNAKECTTLRSDHDPYATYFVLQFPQMFSRMSYSLWKGLVAQF VNLLSTFSWNFIDLFLILISDALTDQFRQLNSRLYAIRGKRNFIVKAMPEWWWTEARMD YNRLASLTRRVDVHMSEIVLLSFSTNLYFICIQLLHSFKVMSNVIQTTYFCFSFGFLLTRT AAVSLYAAAIHDESLLPAPVLYSVSANNYSPEITRFLTQVTTDNISLTGLKFFSVTRSLVL TVAGTIVTYELVLVQFNAVQQIEQSNYTMACEIK >TbGr2 MTISKRMSNNDLNKKSKFTPSKRGIWPFVTKKKLSARTPRINNFVKDPALYNPRFFNRH YFEDPELFQSAIGYPLAVAQYFGLFPLCGVLQSSPTKLKFKIAAPRTLYAYVILIASIFMAS CSILHMFQRLNSETFSVHGGIADATAGAVFYGNNLMAAVLFFRLTRKWSTIQQEWRVL ERYMDKHKNWIRLKLRWKFYTIFWAIFVFAIFEHVLSMVNNNSVDLTSSNMTFEIFLDD YIRKSHAFILSNVKYNFTLGMFIFIISKIATFTWNFNDVFIMLVSTGLAERYKHLNNRIIKT NTGQFSEVDWLELRENYAMLSALVKRVDNNISSIVLLSFANNLYFICLQLLNGLAPSDST LINMIYFFGSFLFLIGRTVAVTLLAARINDQSKLVLPILYNCPTSTYTIETQRLQQQITSDE VALSGLRFFSITRNFMLAVAGAIVTYEVVLLQFNVAIRQ >TbGr3 MSQKITLLSAVRPVTIVNTIFGIGIIEFPRNHPRMALTYLYGLAHIIVYGITVHYAFGEIKYF DISSNVRQFMCKAGLIGNTFLTIATMAIAFQRLNSAMECLNSIVVADNIMETVGIPKDYA RKRRQHLIGIIFLMGTFIVMTLVNASFMFIAGTPIWTRIVLIYATHYPIIIMHLADSTFLYIA SYAGKSFQELNKILSQISKGCNSIFKDGESKVDKIDHYVISCKSDYKEIISIVRESHLHIVRI CRDANVHFGLHTLLSLSFSFVMITGLMYNIYVGLFLSPSSFNKFSHEFISICCWLFFYISKI LALTCICSATVEQSMKTGEIIPSLSMHQEIRCEAQQFLIQMIQNPLEFTARGFCTIDHTLVR QMSGTIMTYLVILIQLSDSSIEIPRSDNNNTLSNNSTH >TbGr4 MSAKFSKKENEQFFRTLFLQLFLLNIYYMILDLLCDIKYFNSSCKVEDRKKLKFKADKTA NTINTWISWTTLPENIKLEIETDIMFRSKKVEPETQNVYSIMESMAGYWRLSGLFPLKLH RPNGVEEYTPSILLIIYSCALFCVPLVTYVIKAVIMNKKENDGTKLSLIEFNVELQEILFAF SDIINELANLQMYMTSLNFHIEFKYRKMKIMTMTFLTVVFLIEGHGLILSLIEYMRGYFY RLNLQLMSLNEKLENVVRKQDTLIFKEMIFSTETIRMIAKLHYGLCSISKRVNKKFVFNVI IMTLTNFTIITMSCYYIFSQLQNDKPTIFFVVICLHRILMNALPLIVFIRVCDWTRYQASFT QRLLNQMELKDSDTVLHDIVYSFTMQMLHQKIDFTANGLFPIDSSHLQTIAGITTTYLVIF IQFSPPIGF >TbGr5 MNKWWSFVKSQVDSQTIKKSSHKVGPKKVVETPYTDLYSALYPIYHISKIFGTLPVTFK RVDYKYRGTTSICNSIYGVFLVIFSATLQLWGLWRDLNLGWQSSTRLNSETAITATCLDV LAVFMLISSIILGAPFRWKFIQGSLDKMVEVDRSLGAFKRRRLHLVTILIASLTFSFLITLSI MDYACWKYAYKLKKPHAPSDKGPLNYVPIYMMYLILTVFEIQYCVLTYNVSVRFQRIN ELIEKQSKGRQIVNYFIKDLALDDPVNRNEKIVAFLDSDLAYFKGKKPDHLWPIIENSDIQ RKIAMSISKLIILYGHMCDAVILLNNSYGITLIIATLSCLIHLIITPYFLYMEINAEIFVWQTI VMQVLWCIFHVLRLLLLVQPCNRTMAEGRKTGVVISQLLSSDWGPEVRKQLEIFSLQILL

417 RPLNFTAYGLFTLDRSLLASVTGVVTTYLVIMIQFQTTDDEQGTKFMLKNATQLLTSVK NNVSLIRSATAIT >TbGr6 MSSNAKKEEPEKDDGKLKGPEAPLYNAISPMFYTFKFFGFAPYKFDGDELVPSITSLGFTI VALIFYTFIFVTFCSRFTFSGFNVNDTAVLITTERGKVVLNYFVLAFSLIETITKRWEFTAT WNKIADFDECLRKKGYVRSESSIRIRMWLLVVYNVVIWFSVNQAGMYAFSEPWYKNVS YLLLYVSSAAALYKYAGIVYCLGQRFKFLNKIAGLDEAAYYKDESQLEKDTKWIEVMF SELMAISESFISTHTWSLFVWIFNLGLHSVSNTYYVLEYCSSGEIDFPYYFVWCLVSWLF AYITQIVLVHACCHSASNEANMMAYLLLDRRRCLAIAGYYKMENLQSTLHLLNRKLKF LAAGCFDVNLPLLQSMIKVTVMYLVILMQLPDERYGTEKDNT >TbGr7 MIVANCLTKQTYILLYYYFKLLGIFPTFFRDETVVASHRAKAYAILLSTVYTFLFIFTMKT RPKFQSTVETPVLIMAKEIMITFMYLLNVCSWLLFAFNQNHLHGIVQHLKKIMALMKKR SMKLNVERIAENVAVYLLFTNLLYLAMFALNERIRWEHIKRTMGTKAVEVAFTVKCFTF PNILCYNITCFYIISLYVVEHAFRLLNKEVRKNTINYLHGQLCDAANLLIRYLSIPVMLAL LVYNVMIFTDFYFIYSTFELTRHWTHFEFIQLINTLSWILRHIVDVYLICSFSEYVKQEVCY SCLFILY >TbGr8 MNLLCMFFNISFDKVKIASCCLRKSIHVSYSVTSCKCNNHTSTEKRVHFIESKKTVIFQML MYLALVGLQINTMCFMYLWNYSNVTFVTKVNDHIFVYTCAISLAFIIPSQWIQQKQMVK LLNQLTDIDCTLLNLRQSINWKILANTIRVLVTFILWVISYTNSYSFHHHAIGSIGFHLPSLI LLNFILQYGSIIEQLAMRFEAVNSILERHFFRQHSITQESIMTIFTIRRFHARLCDIARGVQK CYSLPVTACLATFFYSICNGVHVIISALIRKTFDLYMLNGSIIWCIISIYLLSVLTTNIDVIHQ QKAVLNYLKLRHEKISEDLIFYKIWL >TbGr9 MLLFPFINDYFLLLLSIFKLSPQSNTMCLLRSCDTIFRVFYYSGKVFGLCPFSLNKNQDVQ FSKLGFIYNVTLSIFYMIIGIEMLTIRAHLKFPKKTTLARVMDSFAAFFHCLAVLGTWLQL ALKQNKIRQIVIDLNKSKEFSSNSKKYYFQILFTQITIGIVLYIFIDIATDIVLGYVMQSSKE ELLFVFNFFQIFERVWIIFSANFFSVIRENFYSINKQITKRVLVMKRYYAICGAYDQLKSIL EIFVETFSVSIMLSVIANFIQVNTISYGLYLFLVKEKMWTVHDGITFTILLFWFYINAMGLF MLCNEADFLKKQVFLLKINHFYKSLF >TbGr10 MVDKNATSRSPAGLRLIFYYGKILGFCPFSLDRDNTVQVSRPGLIFNVLLCIFYAAISFFFC VIQERVEVQRETHITRLVGNISLLLQCSSIIAFWLQFGLNQTRLLDIVAQYHLITEMRKKL EFKRNRTLLKILAKYLVILNLLFVVLFVFNLIGWWISDKDKAAAYTFLFFIINLNMMVVF SWCLYITEVRFSDLNEELTGLRKVNENRGKKMMIAGVKIVQQLHDEICNAREKMHNFF AIPILLCILCHISQIMSSVHILYMTVTGTGTWSVNDLWRVTINAILINFSFFELYFAYYFAD NAGNQVSTRFLFPIC >TbGr11 MNLFSKYLPFMTHSSKVFRATNFCSFMKPAFFLCKLFGAHPYKRTRSGLRISKAGNLYN FFMTGFFVLIEFPIQCYSTNIEYYHDSLPSKLQSNCYIIFTFITIVSTTLFRNSRLKVLQSLEQ ISKIVPEKDFYNISKYVHAKDIFCFLFMAGQIFNCKRKSLEGTTDQIFSVYEIVQVYLLDS MYFNCAIVIYLCFKMLNEKLKMIKTKVVSENRHLLRRVYHEHNNEIILKEINEVKKIYQQ ISDFIQKFNSTFGLQILSSFTLGFVEVTFTLYFYLLYKAGMKNININIEKQIWYNYFTTSICY YSVKMVVIACVCQCVKDEARKIGSVLHEALVKITDAEIKEELNIFSLQVLQRSNYLEAKR ITIDATLLTGVSLF

418 >TbGr12 MTMRKSIFLRNRELEFTEPIFIKYIMGLTLYINKLLGMLPYKTEHFKIQKCSTLNCISDIIFV TSTFFSLGILYSCNAGQTPFVTVAGTIQANCFVILGIIIDVAARMFLDSRIYVLQLLNEISIII PSDKKKLSKFCHYKEIFVTVFFLVQISNLIYWNYILTISKLTVFYLTISCYYLDSLYMNCVI ILTNCFEQINCELIQMKKCVLMDEAHLLRREYHEHCNQKLFAKFYEKKNFKKLTKIIKIL NLTFVVHLVCTLFQSFTEITFSLYFYILYRKGYISINLERQLWHYYYFTAFTFYALKFTSIII ACQKAKEEIEKIPWLTNDLVLHTVDKEFRDEVQMFSLQILHQNNVLCMKNVRLDAALL TKVFFMC >TbGr13 MRPCFFLCRVFGCLPYKISKSNFSPSLIGYVHSSVVTATFIVACATVIYHMDISGDLGYTN VPGMLQHNCYTLFACLFTLGTYACSSKRMQLLQDLEVVSSLLPVENFNRNAKFIHAKDI VGFCVLLSLVSNVKGETFYEIVADTAAFFGTLLFFVMDMLYINCVFMINQCFDLITFKLK KLKEMMTTGKVHILRKHYHEQNNPLILMKLKNAIRAYHHTSNILENTNSVFIVQILSTVT FTFIEVTFSLYFYILQNWGHEDLNYNVRRQIWVKYFIASISFYLIKLIAIIFIGQKTKEAAY AVGVQIHDTLMNTTDKQIQYELQLFSLQVLHRDNTLSVKALTIDATLLTAVIKMSYYLF SSCHHESKALKKFELFQIIGGITTYLLILMQFLIAEGSNVKRNEKNI >TbGr14 MLRIIYFFFFWSLSRSWQLFYASDFCSLLRPCFILCRIFGCLPYKITKSNFSPSLNGFIYSTV LTVAFLLSCIIVFYNINISKELEYTNVPGMLQHNCYTISAFIFSTVNYACNMKRMRLLQDI ETVSSMLPASIFEENAKLIHAKDIIGFILLIAITTNLKADKPSEILAKISAMFGTLLICVMDS LYIDCVFTVKKCFDYITLKLDGIKDMMVKDEIHILRKHYHEQNNPLILMKLRNVMKAYH LISNVLQKINSIFAPQILTTVTLTFIEVTFSLYFYLLRNWGNEDITNNVQRQIWITYFVASM SYYSVKMIAIIWVCQKTKEAASAVGVHVHGTLLSTLDKDIKYELQLFSLQVLHRDNTLS AKALTLDATLLTTIAGGITTYLLILIQFLIANSDEEVEESQD >TbGr15 MKSSNFLRSWQLFYASDFCSLLRPCFILCRIFGCLPYKITKSNFSPSLNGFIYSTVLTVAFL LSCIIVFYNINISKELEYTNVPGMLQHNCYTISAFIFSTVNYACNMKRMRLLQDIETVSSM LPASIFEENAKLIHAKDIIGFILLIAITTNLKADKPSEILAKISAMFGTLLICVMDSLYIDCVF TVKKCFDYITLKLDGIKDMMVKDEIHILRKHYHEQNNPLILMKLRNVMKAYHLISNVLQ KINSIFAPQILTTVTLTFIEVTFSLYFYLLRNWGNEDITNNVQRQIWITYFVASMSYYSVK MIAIIWVCQKTKEAASAVGVHVHGTLLSTLDKDIKYELQLFSLQVLHRDNTLSAKALTL DATLLTTIAGGITTYLLILIQFLIANSDEEVEESQD >TbGr16 MSIDKISARLGWIFFKIIGYATYTLKFNPNCDRPKCYYSKLGAIYNVILTVSVSFVMIKAEI DSHYDEDNDENSLIKIIKSAEAVLGISAMVSIHLTYVFAQKSIVHIQNCITEIGLALQMFDH RFEAKSNTRFYLRYYLTQQSVWVVQSVTQQISYDLATSFHLTILSFLIGYFLFQYTMYVK KLSQQLKHLNDLLQALKINHETKLAFYFNSSAKRLDRIRKLHTVIYTLTKKVADFYSFA MLTSTSYLLISQTISVHYLMDTLLTEKDDELFGVFILNNASWIIAQAGPIVNLTNSIDKLSY ESRNTAEIVHELYSMTNNKELKVKLREFSSKLLNQNGDFTAAHLFPINSTLLHSMMTAV TQYIIIIVQFNSSKKDKSKKGQRGG >TbGr17 MKCLPISIRSNSTLYSNEFYTKSDWLYLKFSCLLYKSIGLSSYSIERKMFNSVQKKPVFVIS KRSQIYTILLYLTVVIAQLHIIFKMLMLKSNKYNMLDLILYVLRAGTFLVCVPIYHYRRLS FVKLLNKLVDIEIELSTFKHTIFRPNLKNMCSLLIITISPFILICLNFQNGITIKGINFNIEDNIA ALEEVILYPAHLIFLHSILQYGLITASLVDEFRLINDALISTSFLLYNNLNEFQEKLLALRLV HGRCCELSRNFADWYSLPITICMATSFFFTCAFNYFFVYDTWKKHNQSASFMIFFLYLFS

419 SFYPIFIQTRNVDNLYKQLKEFSAYLNFVDINISAQGLCILDNTLLHSMICAVSTYIFVFM QLNRKLFVDEKLLVNNTQISSSIC >TbGr18 MYVNEFLKIIIPWSFFKALGLAPGHLSFVEMTKAKSIDLKIVYSRYGSAYNFVLASAVITL QSLLWTSFFDQTFPGGPGERDMLIIIYVIMNTITTSNILFHYGLHQKSLVEAANTLVEIDQF LFKHNFNDKKGSQIFGIYLLTLILMVFISVIMLIERLTLDDLNDWWYIYLSHFITGLFILQY SLVIKTIAIRFNVINKSLEELMKNRKLDSFVRTNGFGDHNLYVSVELLRKSHYRIFLVSKS LSRWYSFPITISLGYSLLCMVHGIYFIILNILKYGINEACVSALNELMWLIIVAIPIIDLCRQ VDHLSVELKEKSADLVNELSMTINDENCISQLDEFSLYLYHLKPEFSGSGMFSLDSTMLH SIISATTSYLIICIQFQIE >TbGr19 MKISKSNSEIIASDFPWIAAAWYVFKILGHGAASIVKEPCKLYPSHQLHPKFVPSSFSIFYN FFLMCFLLGVDLNLIVFSYVFELRNVSLVSKIINMIETIFLASTIAILIPYNTIYQSHLCKIFN KIIQLDCEFSDTNLTKSTCKMVVYAFLSNVLWCIALCISGRLNFYDSFLSVLYYIPLMSNL WFPMQYAFLIKILCIKFEIINDCLSNCLDYKRFLPRADKYLKKIINRNPTAQFSSLQSLHSS LCDIANEVDEYYGTPIILCLAQYLYTLTLYGYFIIIYLVQKNKPSATIHCALWFVTVFFPFY LLANRVNEITKEVKKTGTIVQELIVCSKNPTHQNQLINFSNYILHTNLRFTAKGLFSLDGT LVHSILSAVTTYVIIFIQFENKHQSRSCNNNVPSVLNYTNNTSL >TbGr20 MSILLCDTYFISFASYLKLNHFYKVIGLSPFKLKLTSCKVHPNHLIFVNCTKCFYTVITIAI VCTTYQAPFLAMANKVSFINSSNIGLISDIDCQFSFNKFNRIPTDNKIKDRMMRVLNEFVE THFSVVSYFVHDNPVDRFQSFLFTILVVFCILELIVITFTPLPKETTVTWFDIYYTSQYCLPI IVTMTFFMQYSILIKFLQYDLRNLNTILREGHKCRDPVLKDVRNLYSKLCKIALKFHTFY SLPLTVCLLNILISLTTNAYYTVRTLMIGNPSFAMLNNSVGLFVMSTIPLWFLTKFVSQIL NEVIYYASLLLILLTRCR >TbGr21 MKKSLFSSEARFLSFESFLRLNYLYRVAGVSPLKLNVHACKNHPRHLVFSDSICGLLTVI SITAFLILYQLVFLTMVDQVSFVGMSKVSIFSDIIFVVTNLMANVLIGSSLVLKRKIMLKN LNAFIEIHHSIVQLYDGVISAHKCKFILFCVIPIFCTVFLFFVTFNPHPEDDESDWFDVYYS SQFCIPIVMLLIFFMQYTLLLKFIQYEFRTLNMILKRHSSIWNTGRRTFDVDEIKILHIKMC KMAMNIHAFYSLPVTACLLNVLLGITMDTYYIFVTLIQGDIEFGIIVSILGLITIQVVPVLFL TTFVSQILNEVKRTGIIIQEIIVTTNNQQYISQLKEFSFYVLHIDLKFTAQGLFPLDGTLLHS MLSAVTTYVVIFIQFDSGNGSKNVNNGTQ >TbGr22 MSTQRINLKEFGYIYICAFIFRAMGLLPCSVKKVKHTGHVLTLNTPVFKNSKGAIVYLVF LLIVSYVGMLRVLMLYFDGNVREVSDLGFIADVTDIVLMLLSFTIILTYYSLHHRNFIRMI NQMVQLEYKFRNLRHQTDTYYILLVFFFFVHIFVWITLLPIYLIYFDHDVSIGCYYLIFFII QWFILQYSMFLKVLIKHLKTINSMIKHSSIGNPSRNVIEFVQFQCNETFANEIMVNLRQLY ATCVAVTQSVSNFYSFPILTAICFFLFSLSANFFGFMSIVKTEGDKIAMMFFLIIWIIMELTP IYHLTQSVNNILSEVKQTGKILQELIISTNNQQHLAQLKEFSYYVLHVDLKFTAHGLFSLE GSLLHSMLSAVTTYIVIFIQFDSRVECDT >TbGr23 MCKKQVSDFWRTFYTIIKLTGLLPCDYKKSAKSRFPTFYYSNLGNLYNILLVIFTLLIEFD LFYYLSNLQLSHESSLVFTTNTVRLVSGLVGFLVILSTITWNQQSLVDILNDIVRLDGQLK INHRWKFIVLLIWSYFISIIAIFIYSYLSSPPTVRIIVLLGSHICFLIIYCYVLQYALVVKVLT MQFAIINENLQQISNNKIQVLSDACSNFKVKLTFDPDIMKLRLLHEEMIGVAERVNDYFS

420 FPILIVMVVTLYNFSCNIYFLCETYFLMVCLGETIRTNNSTFDYIDFSSYIIHTDLKFTAHG FFPLDETFLNSVIRITHLKK >TbGr24 MFANPFKPSTFVFYLIFKIAGLMPFSYIRNKNGQILTTRCSLIGVCYNFILLILVIVSEIDFA NNLSKYDMMNMTSLVSASISVHAITSIIGSLVILLTFCLRQKKFITILNAIAKVDHRIAFKQ KTEFHWILNLYSFCVLSMIMYNYIVQGEQISLLHFFALYGGYITICYAVIQYAIILKMFRD RFTTVNQKLRDVSRKIEFKNFTCLCNSIGVPLLLTDLKLQKLREVHEELIGAAESVNEFY GFSMLISTGVILYNIASNTFFLYVLSSNVWSEFILTIFYCLYKFVPLWILATFVGQTLKEVK RTGRIVQQLLISTNNQKYLQQLKEFSSYVLHIDLKFTAQEFFPLDETLLHSILSAVTTYVVI FIQFDKGGSGTKESQNSTE >TbGr25 MSSQSVTNCSRLTSQLLYLLLKAIGLAPFSYTQTKSQLPSVCYSQAGVIYNIILIIVLLILKL ELLADLSANKMLNPTSLMVTSNVIHIVGGFISSSTILLAVSWNQRKLVKILNEILKIDHHL NIDKDTKFVHTVVIGQFVFVTSLFIYNLSVLLEMISFWRLLGGMYVTIVIIISYIMNYALIV KMIQIRLEAINERLKEFSKKPRTRQVVVLEWKVVSLDGDLQKLRNLHEDLIIAAEGANQF FSFPILVGLAVLLYNFASNVYYLYISTLEMNYHKFIITVFWNVYKMIPLWMLTRFVGQVL REIRKTGRIIQQLIILTMSSDYLEQLKEFSSYILHIDLSFTAEDFFPLDGTLIHSMFSAVATY VVIFVQFDNRSQK >TbGr26 MIQINRNNRVSNFETFALSYNFFKILGLSPLAVKYSNTKRRIEFVNSFSGSMIAILLTVVNS IFQTVLMSLSSHVSDSIMSSISFYSFCSILLLDIATTLIIGIQILTSKKNFLDASTKLIKLHYEI EELNLKLKQQSQTYFYIILLILYYIIMFIYNIYSSVPNDPKTTLLDKFYLNQFWIAFTWQIF YTMQYAILVKQIHYEFQDLNYGLNETNILDSETLSKIKKRHRLHVNLCEVVELVSKFYSF PIIGCLLTFLIANTFNVYSICDEIRKGENCSIRLVIQYCLFFAAQILPVTFLTQSVTDTQREV FMNTIIKEIFLDK >TbGr27 MPAPKWNNYRILDTVLLLLLFKFCGLATCSFLNFNFLRTGKAPTFIPSRITQIYNICLIVIVI SILIYLFFGMQNMNLLILNKVTTVTMYGELASSFFAALAVLLLYTVRQKSFVDIITRLMK SDLGLTTLTKIQIVPYRYNCPLVMLHLSIYVVAIFLSQEIFSDGLFGIAYQVSYLILQLFMY QYEFFVRNIQTKFKIINLTLENYFIEKTVDLGFNQYRINIGKELMLNLRQTHLTLCEIAKD VADFYAFAFICCAASTLYGTTMGFFNLYKTIISADVIALLFLNNILYIWIYFFPIWLLSMTV TNCTFEVCNFEKSYKYNQGNCDEFYINNSFES >TbGr28 MVKRNEFWDLVIYASYFMLKIIGMAPFSYTRKNSDRSINFQVSVTGIAYNLVFLIYLIISGI SIGQHMPEIKTNYYSSVSYASNFIYMICSLISCSMIVISTCLNQKKFIDILKTLVAIDQSFPS KIKYTKFHITVIGYSIAIIISLYYNFLVNDQIFSINHVTTLSLTFVIFNAFMLEYVMIMEFIHV KFQTINDKLLEMTDKISTLSFVAFYDERKQLHVNVFPIIKFRTFYKNMIKVSEQVNNYFS APILPCLVTALYCLTNNVYTLLNLLGTLNVDNCIACTIWIICMVFPAWRLTHNVNRILKE VFIY >TbGr29 MYLKHSLSLLSYVVYLMYKLLGLAPFTYYQLSENEVLNFKFSYLGTVYNMILLTSAVVT HISCLIFIPNIKSKSLTYVSNICFICNTLVSYAMIVTKFIIKQRKIVNTLNKLMVIDHHLSKN GRNYHIELVVLCHFISTVMCLCFTFETDSKFYPQGFVIPLYALFLIIYCYTLQYAMILRFIIR NFKSINETLLNFQNSLYCNINTRCNFKFDITTNNTWVDNLKCQYKDLIKLSENINDFYSVP ILAICTSSLYSLTYSTYFFYISNVNCNIKQIFESVFWIASILFPIWMLTSAVNHILNEV >TbGr30

421 MNFKLISNSSLSYLLYLIYKLFGLAPFSYCHNQVNELETFKCSFFGTAYNIIFIIYIASSHIFC VYYIPEIRSKSVTELTYVSNYLSTWSNLVCGVLIILTVSLKQKTIVKILNDLATKQHHFPK KIDNHNIAVIILAYFIFMIVSFSYCFSTDNDLYPLVYLIPLYAAEFMTNFYTLQYAMILRFF IADFKSINKNILEFKKLSMFEVHSRGPEKLNFITNIVRIKDLQLLHKNLIQISEKINDFYSVP ILAVLAKTLYSLTCWIYNVYLMILVPDYEILFEFVLWIIIIMFPVWMLTSVVDHILKEV >TbGr31 MNIKIILNPLLFKIIYLIHKIFGLAPFSYSQNQNSDLAIFKFSFFGTAYNIIFIIFMTFSQLFCIF YISETKSKTPTQLTYISSYVAVGSNIIFSVTIILTINLKQKTIINILNNLIINERHFPKKTDNPH IELTVLGFFFFMVATFSYSYFTDRDIFPFFYLIPLYTVHFTFNFYTLQYAMILIFITKFKSIN DNILESEKLSIFKVSSEDAVTLDFMTNNVRIENMRLLYQNLVELSGKLNDFYSTPILVILA RNLYSLTSGIYYLYLMILIPDFEQLFHCILWIALMMFPIWILTNVVEDISKEVRYMLFPFFY NLTAKYFKRAKFIERVKMRFAHSKQILFGTF >TbGr32 MRMDLKHFFNNSLSYLIYYLQKLLGLAPFYYRQNQDGECFTFKFSVLGIVSNLILLTYVL ITHISCLFHIPKIKSEYLTALTYVSSYFFVWNNLICSVIMILTVCIKQKTIVNTLNELVATER YFNMTDSNFRIKLIVLCYFMFMTVSYYFMYPTDRKAFPPVYLFPLSGVLFATNGYILQY AMVLKFIVKKFESINYSLLQYKNSSDCNISSCSIFKFDHKKINIWIQNLRLQYKDLIELSKN INNYYSLPILVILARTLYSLTHYMYYIFLMMSINDYGILYELSFWIVTMLLPIWILTSAVSH ILNEVRYLLKINQKNRHLLNKI >TbGr33 MNLKIILNSLSYLVYLIHKLIGLAPFTYNKNQRNKLSTFKFSFLGILYNVILVIYITYAHISC LFTISEMKSKSLTAVIYVSNYFFVWNNLICIVTIFLSVIIKKRTIVKILNKLVAMDRYSPTVI HNYNIELIILFHVIIMITCFCFTFNVDRVSFRSEDVSALYSVNIITNSYILQYAMILKFVIIKF KAINNTLLEYQKLSNLEVSTRGISTPGSTTDSIDIKNLRLLHKNLIELSESINNYYSMPILVI LASSLYSLTNGTYLFYLTIEIPDAQQLDWISFVAYDHFVTDLDVNQQCKSHFKGGTILQN KNY >TbGr34 MFVITRLSHLKMNFKIGRKSLSYLVYLIHKLFGLVPFIYDQNPHEKSLTFKYSFLGIIYNMI LLVILVYTLISCLFSIPEIKSKSFTKVTYVSNYCFVWNNLACIISIILTVSVKTKTIVDTLNK LVAMDHLFPKTIYNHHVEMIVLCHVTTMIICFSLTFDVSGVSIHLVDNLSTYAVYIIINNY TLQYAMVLRFVIRKFKTINETLLKCQRSSNLDISKRKFVTFNFITDIKNLRLVYKDLIELSE KLSDYYSIPILIILASNLYSLTNGTYFLCLMIATSNVRLQFGTCLWIMITLLPIWMLTDAVN HISKEVRRCTLDAIRKLRGLNVLFLQKLSLLHFIYFTL >TbOrco MMKGKYQGLVVDLMPNIRLMQYTGQFMFNYYNDNMKMMHLIFCCINLALILIQFAFC LVNLVFDKEDVDELTANTITMLFFTHSVVKIIYFALRSKLFYRTLAIWNNPNSHPLFAES NARFHAIAVTKMRRLLFCVGAATMLSVAAWTGITFVGESEKRLHNSETNSTDIIDIPRLP VKSYYPYDASEGMAYVLTFVYQFYWLCITMIDANCLDVLFCSWLLFACEQLQHLKHIL KPLMELSATLDTVVPNSGELFKAGSAEHLKDTELAPPPPPPDVPAGDNMLDLDLRGIYS NRQDFTATFRPTAGMQFNGGVGPNGLTKKQEMLVRSAIKYWVERHKHVVRLVTAIGD AYGVALLLHMLTTTITLTLLAYQATKINTVDVYAASVIGYLLYTLGQVFLFCIFGNRLIE ESSSVMEAAYSCHWYDGSEEAKTFVQIVCQQCQKAMSISGAKFFTVSLDLFASVLGAV VTYFMVLVQLK >TbOr1 MWKSPDHFWTKPKRNSNDKKNSIKILRPPKFLIQLVGLWPYFPNYRQYYLVMSYLTLFL ASLLIDLVMNFKNFPEIILSVYKTIASCQGFVRIIVLKKHIYEIKIFFEFIKVEFNPSNYEHQ

422 NEKNIVMFYNLKAKKFTRFVSIAPFISFFMACVNPMFTQAIPILINKQKFNYVLPITLAFYT PHSWTSFISLYITQTLFLVFLMFSVMTMDKILILIVYYLCCRMTILSERIRNLNVFKEADDL ELFKIIREHQKICRLAYKLDKIMNIWSTFYIMFVGGQICGILYIFNVLMQNRVYVIAIPSIY FILLECIKLYSYCHQGELLKNHSLKMSDTLCETNWYTFPMNSQKKIVMIMLRGQKAFRL SAGRIRLFNFSQYIDTLKTATAYYSFLRNTMHSQNNFSMINESYKSS >TbOr2 MRFRTKSASNRAAFKKAASILPWNRNLLNVMGVWPKTPNIIRYGTAILYYIYCLGLIYLS FFLALGSFRKVLRTLSEIITFTQVFYRLVTLKRFNSEFGEMLENIKRELDIKQYRNEEEMK SVLWYNRVCKFFCKLLVYCLGCTVLLYYFKPLLLQLGNKLTAPKNVTVPLTYELPQRF YSFYKVDNIRTYALTYIFQMPFAFINNFTQASMDCLTLTVTYHLCGQLAVLSLRIEEVGH KPIDETHSLAHHINKHQQLLRTGGILAKILESCLLVHLVGASVLICILIYNILFMLAIGQKT GLLTFFVYGSLNVARLFTHCHAGEYLIHESLGLCEAYYNCKWYELTIQEQQSMIMCLKR SQKPMKLLAGKFGTMSLTSFGEVMKTAMAYLSVLRSLI >TbOr3 FQILKNEKWVLKFNKIYLSILGLWPESPNNLLFNVLFMYYGYCLILVYLTFYLALGNFRK VLRSVSEIITFTQIFYRLITLRKFNREFGEILDDMKNDFKIRSEKTNEENRVISNYFRLCKLF SLILVTFLGTTVVMYFFKPIITQVGPSMKKELCNQIRKNENVIFNLKLKILGIFQPGKSHAI FKYELPQGFYNFFEVNSMIRYIVSYLLQFPFAMMNNFTQSATDCLTLTIVYHLCGQLFVL SSRVRRLDYRSQADKNSIRHYIKIHQKLLGIGTKIQRSLQSCLLIHLVGASLLICILVYNVL MSIARGHTMILLTFFVYGFLNILRLYAHCLVGEFLIYESTKVYQAFYEFNWLSFPIRDRKL LIISIRRAQKPLYLMAGKFGSMSLTSFAEVMKTALAYLSVLRSVV >TbOr4 MIIQKDFDKAAKIFPWSKWFLLPMGLWPLASSSTIFFLSFGYFMYHMTLDFIDLYFSLQS LALLVTNLTETLPFTTIISRLWTLHSHNKTLKILISTSLKDFNITKYQNEDEKATFMSYNFI AKLYVKFMIVEILFTGTSYYLKPLGQLGEILAPGNETVFNETGSIVFPMPYRFFLVYPIEN FQTYLMTYASQIPFLIIITFSTAGGHCIMITLIFHLCGQMAILNIRFRALDNIRGPEKSRELRI LVKEHIKLLEMGRAIDDSFSSAILVDLICDTIGLCILGFHIITILAEGGGAGLVTFGVFMSLT LFVMYIMCSAGEQLVTESSKITESLYKINWVDMSKEDASCLLFCLLRSQKHLCLTSGGF GTFYLSTMTEVIKTSMGYLSILRKFMT >TbOr5 MFKGQRDLNVAKRIFLWNKLSLNVLGLWPPSPSDFRFFLSFGFFGYHMILEYLDLFIFIG DLEHVIMNLTENLAFTPIVSRILMTRINLSKIRMVVSESLLDYKADKFRNEDERNVFLPY MATAKTFVKLMIGNVGMTASSYFIKPIMGRLGADPEQNITLGYELPYRFYMLYEINDTK TYVWTYSSQVPFVLISFFGTTAVDCLLVILVYHVSGQMAILASRIRQLPTNDTPENLEEIY KVIRLHIRLMKMGEAIEKSFSVTLLVYLLCATIAVCMLGYQILTNLDKDIAILVTFMIFMC LILLLLYAQCTVGESLITESDKVYKALYECLWYQMPCSHSRNLIFCMMRAKKPLHLSAG DFGTFYLSTMTDVIRTAMGYLSVLRNFL >TbOr6 IYTTVLFTKLGYNLKNEVILNHVVFGTYTVLAFVKTSCFWYYSDRVSQILNSVSNDYLN QNNVQYQEIIKLSRFEKTRTINFVRIFAIYIPVASFQLTTLLSFYIEGYQLQNCMPSSLADS YIFVLFTRCVQIYFIVAGNVGHDLFFVMLAMYLCDQLYILGEKYSNLTKNSSEDECLLRL CKLIDQQCHLIFILKNIEFSFCTVLSLQNIFGLLVLYIAGNEVSEENKIVVLLRSTLFISIIFC QLFLYHWTGSNLESSLDRMTQAIYFSEWYKTPWKIWKNFCLVMMETNHPRFTNGYIFE MNIITCRTLAHFLSSLKILIGYLCQCSKMLLAI >TbOr7

423 MDSKERNRKRKELRSQLFSPAFLFEFNLYKIPCWPCGTWPLERETIFKDVKFLLAFLTQV STTTILFAEVYFACAETEVLIDYIGLGSCSMLAFSKTATFWYYSDKIFESFLWMSEDHAK TTDPNYREIMTMYAGKARRIFKIQMTATYFSALSFPVNVIVLELLGIVKYVLPMQTVCTV SNVTGYAYPPAFLEQCWHIYFTATGNIGHDLIFIGMVMHVCGQLHILANEYSKLTDNSK EKEYWSKFGELIERHRHLYKMFDNMSQSFSKMLMIQLTISVGLLCILGIQILVALMENNT GLFIKHTLFTAVVMFQLYLYAWAGDYLELNYDRMTNAVYASEWYKIPTKTLKNMIFVL MRTNRPTRFIAGYIYEINMITFAAMLKMMMSMFSIVRLMFIEEE >TbOr8 MNLTSKIELKFYELSLQVCGTWPLEKLCFFRNIKFIFAVTAQLLVIPMSFMKIVDSCVENE TLIDCIYLSTCSSFAMVKTTNFWLYSKKISEHVRSVAVGKNEFFHEGEIQNTDDYKKAK KLFLFQTVLTYIPSNIILVKSMLNIQNKHNSTSWTHFPIVVLKCVQMTFVSIGNIGHDLFYI FLVLHICSQIRILEYKYSKLVMLPNEKQCKRKLSMLIDEQQRLLTLAKNLDISFSKILVVQ YMLSLMVFYIFGTKSLLTHLPENTAGLTINQTLLTSVFLYQIVIYSLSNHYLEHNFKRIKN AVYFSKWYTMPPKLLKNVVFILMRTNHPTKIIIGSFFGTDLIDFAVLKMLTAMFYMARC TH >TbOr9 MRAIFQSSKKQKIKNHEKEDDIYNVGISYIRKMLQLSGGCPAAAGDGFLFYLKKYLALLI TFLFLSTYSFTIIAGVMISQNDIFALLVDLVMISFVFILMTILVIILNRANIITQFFQNMVTDF EELEADSRQIMLDYHKMGRKLTITSTGCICITTVVWNLLPLLDNPVQLGNKTVQNFAQT NEYFLFDSNEWPWPLFVRLHQCISIIQCGIIMGNMCCIVVGFTMHNCALYQILFKQFTDF CVTDKEQKALKFLKFAIKKHSDSMLLTMTIEKAVSPIVFFEMMRCIIATCVCGFIFILDDG EDLFQWFMKLFSILGAFTMTFVFSYTGEQLVTNCDEMIAVLYKFKWYLMNKSNCKNIA FMLHISQKPKYLTAGKLAPLNLFTFSDIIKRTVALLNLFRTMIE >TbOr10 MRKKYDDLLQYNPNYLNDLKYAFNINRCLLYLTACWPIGTRVFKTIKRIVNFIQICFIIIFI LGYVFDLILNVTTIGEYISKGGMLLVFIGYFIKYLVTVMQQSIIWSCYFQMKEDWKYFW TGSYEIMRKHASIGRNITIFYCVIIMVTLALWNVIPLNKPAVIVDGEAYHNLPFVSEYIFFE PRKYPYYPYVFVWQFYTAVLIVVSVCHNSTAAAFAIHGCAQYEILLIILRDISENTKHDV DANTKLVYVIKKHNQILRFIAQSDRAFTWIFFTEWIACVNSMCTACYFCLVEPLTLMSLL RNIGLFTYATNTFFLICFTGQQLTEKSSQFGKLFLTTKWYNLTKEKLFIFSLINVYGQIPAY ITAGKMIKLTMETYTDSMKRTLAFFNVMKKTL >TbOr11 MASTAVQTSSPKFEKRWSIKDRYDRFIAFNKNYKKDVDFAFRTNRFLLYWTASWPMKT PKLKLVKILANIIQTTFLIVFLIGCFIELFTNRDVTTSEFINKAGTLAIFVGYFGKYVITASSG SQINVLINELTHDWKYIWESGYEIMRSNALFGRKLSNYFLYYLAGTFACWHVNALNQK ALIKDGVKYHNLGYQCDYIFFNVRVYPYFPIVYIYQLYSVTVTLASILHNCTGAAFILHA CGQLEILMQIVKEVCNDNDDPDVQIIKLSYVVEKHNKIMRYMKLFNNVFTWIYSTEWIA CISTLCLACYLFVTEPLNPPNLMRDISLFFVGVNNFLLICLIGERFSQSVSKMLSVEVDCII GEVLYNLQCEKFGYLFVITDWYNLKKNHIMKLAMINKFGQIPAFTTAAKIIPLTMETFAD VMKRTLAFFNVLKTTL >TbOr12 MSTNIRIIRNPTYAKDYESKFNKWLFYCAAIWPYKNHYRIKVKILQRIMEMIHMMMILNI AIPLMIGSYRTFNCTENSIQIFKGLVPCVICSCLVVKYIVNVENNHAIYTCIEQVDLDWQY IFENSRETMLKYAERSRRLFKVLGFIGLSTPILWQFVSLLKKPVVIENVAYHELQLIADYV YFDPRVEPYYYYICLFQYAGQCAGVISLISACQPITFIFHTCGQYDIVAAMINKLKLGDSE TVKETRRVLTLILKKQHSAMRLSSDIEKAYHVTFLMEWLIGILGSCFSIYLIVQEEYTLMG

424 LVRNLSSFCLLTCTFSILCFCGDQITLKYLDCGNVVYMSKWYALPKKDSIFPVMMSGIFR ISKQLTAGKFKKLSVRTYADEMKRSYTTFGLLENLS >TbOr13 MEGVSLSYTNNPHCERDIKYSFDSNKILYYITVVWHVKNESLFRRLAMFLVQIFFNICTF FFVLGFALDAIINVHNGELRMKMFLNGILFVGYHIKYIILIVQSRNIRLCYEQLKCLWENI FSGSRKHMMSYATRGRKMSIFSIGLVYIVVMSWQAVALVEKVRVVDNVTIHNLPYPSY YLLFDSHLAPYYYCMHSYHILVQILTGAVVSCTCTPAVFAIHVSGQYDLLIHFVKMTLN ANDSPVKFQSKLQMIIKKHRDLIGYTDLIEQTFKWVFFIEWMNCIAGVCLCCYCMLTEST AVTYFKNFVTMLFFMALFFIPCSVGDELTEKNVFLSESGYLGDWYNFRVQDVFCFYMIN AKKSSPNKLTAGKMIILSMETYSEVLKKAMAFYSVAKNAVVNK >TbOr14 MGIEVNVIENASYNDDIEYAFKYNRWLLCSTASWPDVNSSILKVCFKILNMVQIAIMVV AIAPYFISMFSTGVEIQELVEYLGLLNVCLGIFIKYIIILLYKKTISACIHQMYLDWRYIFEH SRETMIGYAKVGRFFLIVLTTMWLSSWFSWHLVVWLKEPLIVNNVTLHYLPYDAYFEN VDAREAPFCYYICALQFLCDGVLCLTACNICLVISLAMHICGQYAILESIVLNIANGEIND RQLNICLARVCRKHVTSLSFLNVVLPQGGRCNFVNKWMVSHLEYAFSWIFFIEWFNCIL GVCLTVYVMIKQPQTFNDFSKNMVSVLSFISTFLIVCYIGEALNTNSLRLGQSAYMSKW YKLTPKQNFYVKMINRVHQFRNEMTIGKMGVLSMELFSNVLQKLLAFVSCIKTIL >TbOr15 MNNFFHHATHLLFCSRQVSLSRSFLKEVVQFHYKLMISIKVVENPTYKDDFHKAFKYNQ WLLFAISIWPSHKRPSSFEKVIIQIITFIDTSLMMAFMVPIFLKLTFLQQDLRQMMDDNGV LAVFMGYFGYFLIFKTHNHIIHDFIEQAYLDWKYLFQGSRDIMIERAKISKIILIVIISVWLT TWTSWLILALATKSYCVDDKIRHILPYPAYYIFFDGALVPYYYFIFVLQAIGNLVFVMVIC NISSIVTFALHICGQYEILSSIIINLGDNKGDYNDANIWLARIVKKHYTLTKLTSYLEDTFT WIFFIVWFTGLSGLYTCSYYLVVDEQNISALMRNFTSINSLLCSLFVICYIGEKLTQECLRL SEAAYMSKWYNLSNRDANSVYIINSIHNFPKQLSAGKLNLLTMQTFTDVVKKTMAVYS CFRTLS >TbOr16 MGTKIEKIEKRYLEADYDYAVGVSRLLLRFTGSWPEDHMTKRNSIKYLGSKFLNLVTFV TMVLFIVPIFLEIIFVKRAIHLFLLDISFLIYFGSSLTKFILFHILKYRMYPLLQEIHFDWDRA TPEERKTMFISIGTGRALMLVCVTFMYVGIVGWDVKGLTEKRTVVDNQTYYPLTFPGD CVWINVHRPPYYVYLATIQTFGCYFGLTIMMSTLSMAVCLITHVGGQYEILNNEIKSFEIT RKTETEYMRNVFPRFVKRHMQLIRFSNMADRIYDSICFTEWINCVNGICICAIQLVTVILG AEYLSLFGLHMMG >TbOr17 MEIKETIRNKYYKADYKYAVGVCKNLLWITGSWPEDHVSKLSFKYISAKLLTLTTFLVM VIFLTPMVLEMIFDKIPIYQFLLSCSFIVWFGSAFTKFILFYILKYRLVPLLKTMRDDWNA ATPEHRQIMIQSIATGRTLMIVCTVFMYSGAFGWNYKGATQDRLIVDNQTYYMLTYPG DYILADIHRSPYYFYMAILQPLGCFFGLTMMINTLSMAVCLITHLTGQYEVLNYEIESFEI TEVTEKDFVQDFLPRIVRRHVQLMRFLSSLGTIYDCICVPEYMNCVAGIFTSAILLVTNG MTIVNVINYVCIGMVSSSCMFFLHCYIGENLLEQSLNVGKTIGLSNWYNLQSEQMKSCC MIIFISQHSQQLRAIKIIILSMNTFCEVMNRAFAFISLLKTMMS >TbOr18 MDDTKGELINPQYESDYDYVVKLSRLFLRFTGSWPFDHVEKFKLPNVLSKLSNILTSVIM AILILPICIDVCIDTGPIYTRLLKVTFIVFFILGFIKFFIFHMIKFKLIQLIEEIREDWKAATPEH RTMMLDNLKAGRILLFSCVCSVYAGVIGWDYNGLTGERTIVNNQTYYPLTFFGEMIFIDI

425 HTPPYYVYLAIAQTLGCFYSLTIMMSTFSMCLCLITHVCGQYKILNHDINTFEITTENEKQ FMKYLFPKFIDRHVRLNRFSTLTSDLYDSICFTEWFGCVNSLCMVTINLVTLKLTAVVIF NYMIVGVLNSLLMFFIYCYLGELLQEQGFKVGRTMSATNWYNILHPRMRDVSMTIMTS QHPNQLRTIQIIVLSMSTFCDVVNRSFAFMSLLKQTVTSPE >TbOr19 MENKTEIYKNPHYELDYKYVVGVCRFVLRLSGSWPYDHVKKFTLLYMGSKFLNVITFII MILLTVPIAVEVCFVKEPTYLLLLNTTFVLYFSLGLVKFIIFHMVKFKLIPLLEELSADWK RATPTNRKIMMNSMKTGRSLLFACVIFMYAGVVGWDVKGLTGPRIENENLTYYPLTFS GNLILIDIHSPPWYSLFATAQTLGCFYCLTIMMSTFSLCLCLITHVCGQYRVLNHEIKTFEI TPENEKEFMKHFFPKFVNRHNHLNRFSTTTNQLYDSICFTEWLNCVNTICMCAINLVTM KMTFAVIFNYLIVGMINSSFMFFIYCYLGEQLQEQGFRVGEAISSTSWYNIKHPQMRGCS MMILTAQHPKQLRCMKLIILSMSTFCDVVNRAFAFISLLKQSVSSPT >TbOr20 MKLFGPKENETKKTKNPHYEADYKYVIGVSRFLLRLTGSWPYDHVKKFTLFYMGSKLC NLITIFIMVTLTVPIIVGVCVDKGPTYIWLLNITFVIYFVLGLAKFFIFHMVKSKLVPLIEEI YLAWKEVIPEHRQIMIQTIKTGRYFLFAYLTSMYVGVVGWNTSGLTAERRIIDNQTYYPL TYSGELVFIDIHRPPWYPYFAAVQPLGCFFCLTIICNTSSMCVCLVTYVCGQYKVLNHEI KTFEITTDNQIKYMKHFFPKFVNKHVHLNRFSTVANQLYDSIYFTEWLNCINAICMCAIH IVTNKVNFFVVFNYIIVGLTNSAFMFFIFCYLGEELQEQGFKVGDAVSSTNWYNVPSRM GDFSLISLMGQHPKQLRTIKLIVLSMNTFCSVVNRAFAFISLLKQTVSPV >TbOr21 MKKKSVWLRNPHYINDCNYAVGACQFLLRLTGSWPDDHVKKSTLVYMGSKFLNFITFII MVLLILPVFVSVFVKTQPTYMWLLNITFVIYFIIGFAKFIIFHTVKSELIPLIEEVYSDWNEV VPKYRQIMTKSIQTGRYFLFAYVFSMYAGVVGWNVIGLTAERHVIHNQTYYPLTYPGEL IWIDVHRPPYYAYLAAIQFIGCFYCLTIILSTSSMCICLITHVCGQYKVLNYEIQTFVITTKN EMQYMKYLFPSFVNKHVHLNRFSTITNQLYNGIYFTEWLNCVNTICMCAINLVMHKINF LVVFNYVIVGLINSAFMFFIFCYFGEQLQEQGLQVSGSISSSNWYNVPHTRLRGCLLMML MGQHPKQIRTIKLIVLSMSCFCNVVNRAFAFLSLLKRTVSET >TbOr22 MWNQNYLVDADFVVVVARRLLTPIGIWPRYGNDSSFARFRTYVHVILVFCLMLFLLVP HLTYTFFDAEDLKKLMKVIAAQVFSSLGVIKFWTLIINKQDIRFCLEEMERNYRDVESEE DRAIMVKNAKIGRMFTVMYLSLSYGGALPYHIIMPLVAPRVPKSTGNETMIPLPYPCDY FFFIPEDTPLYQFVFVLQIFISSIILSVNCGVYSMIASMVMHCCSLFDVVAKQMEFLVNED PKHLLPRLKKVVKFHMQAIEYSEKIEKALNLVFLCEMLGSTVILCFLEYGIIIDWDENNM LGTGTYVVLMTSIYVNVFILSFVGDRLKEESLRVGESAYNLDWSTYPKNVTMILIYVIAR TQKPCVLTAAKMFDISLQGFTEVVKTSAAYLNFLRALEL >TbOr23 MQANIKYSISWLKYLLIPLGMWPNSNKKMKYFDLCCIIFIYFMIISMFMTVYLHLTFEAD NSFVKIENTGPMSFMVMNILKMTSLLLQVHNIKHFIKIIKNDWISMNLPHEQYIMLNGAK KSRYICFGSIAFMLSGLLMYHCFIPISVFIIHYKSQKYVNPLCYPVYSTFFDITKNSTLITLFI LQNVIALIESNIGVSMICLIIVFVTHVCSQLEIVMLQLNHLLDDYTKNPKHIQNKISATIKL HQKTTRFIQKIENIFTLPCLIEFMGSTFNLLTLCYYVLRELDEKALFKLSTYSMLTLSFLFN VYVICYIGEVLKQKGYEVYEAVYEVKWYNISRNHSMYLLFILMRTKQPIELTAGKILGLS NMSFMKVVKSTAVYFNASRTMIY >TbOr24

426 MSLDHLSKTTFVNANLKRDLKYALNWNRYFLHPIGIWTGDKQSFLKSICYKLIVLTIYFL LFFLLVVVGLYALVENPSTRMTFVGPMSFCVMAILKYHFLILRTMDIKKLFLHVKDDWN NLLTENDRILMMKGSKSGRSLGFLSAGFMYFAWIFFNNIKPISQLIVNIIKNENVIPLSYPV YSKIVDASESPAYEIVFLFESLFGYLQGTITVALCSLAAVLVMHACSQLNVVMRQMNM MLQKAENDIPVYYNIVAVIRKHLRILRFITEIEKILSLASFVECLGCTCVICLLGFYILSGLE DENFVALFSFAILFLSLTYNIFIFCYIGEKLKAQCENIGIVCYNMKWYVLSNKEAIDLLFII RISQNPAKITAGKFIILCHSSFTNDRNIINENLHLMEDTLSRKSTVTLKISDEQLIKLQSNRV NVL >TbOr25 MSLDHLSKTTFVNANLKRDLKYALNWNRYFLHPIGIWTGDKQSFLKSICYKLIVLTIYFL LFFLLVVVGLYALVENPSTRMTFVGPMSFCVMAILKYHLLTVRTKDVRKLFSHVKEDW TSLSSEDDRIIMLKGARSGRKLGFMSAVFMYSGGLFFHGVIPISVLIVNLNKNISVKPLTY PVYSKIMDASKSPTYEMVFLIQTLIGYLEYNITIALCSLAAVFVMHSCSQLNVVMRQMN VMLEERTHKVSFHQKLVAVIKKHHRTLRFVAKIEGVLNLACLVECLGCSCNICILGFYFL KGLEQGSVIKMITFVILLASFTFNIFIFCYIGELIKAQCENIGVVCYNIRWYILSGKKALDLL FIIRISQTPAKITAGKFINLCLSSFTNVIKSAMAYLNVLRAMAVDGFEFDVKEL >TbOr26 MFHKQLPRTKFVNKNLKSDLEYALSWNSYFLLPIGIWTNGQPSLLKKICNKLLVFTIYFT LFFFLIVIGLYALVENPLSLLSLLGPLSFCMMAILKYHFLILRTMNIKKLFLHVEQDWDHY LNQENKLIMIKGAKSGRNLGFISAIFMYSGGLFFYIILPLSYLIVHLIKHEYVKPLTYPVYS KIVDASKSPTYEIIFLLEWLFGYWECTITIALCSLAAVLVMHSCSQLNVVMRQINMMLQK EDDDFSIHQKLIAIIKKHHRTLRFISEIENILNLACLVECLGCTCNICLLGYYFISGLKNENV FTLFTFVTIFLSFTFNIFILCYIGEKLKAQCENVGVACCDMNWYTLSNKLALDLLFIIRISQ RPSKITAGKFIILCISSFTIVSTNFTINAINTLILYNVDFQVMKSAMAYLNVLRAML >TbOr27 MFQKQLSKSTFVNRNLKSDIEYALSWNNYLLRPIGIWTDVRPSYVKRICNKLLVFTIYFT LFFFLTVIGLYALIENPLSRITLVGPMSFCMMAIMKYHLLIVRMTDVKKLFFHVKDDWT NLSTDDDRVIMMNGAKFGRKLGFLSAAFMYSGMLFFNFIIPVSYLVVNLFNHQYVKPLT YPVYSKIFDANKSPTYELLFVTQLIFGYWEVTITIASCCLAAVLVVHSCSQLNVVMRQM NLMFKYKEDEMTIYHKLVAIIRKHLRVLRYISEIEKFLNLACLVECLGSTFDICLLGYYVL SELKDENFLTLFTFFILLLSFTFNIFIYCYIGEKVKSQCEYIGIVCYNMKWYILCSKQALNL LFIIRISQRPAKITAGKFVILCLSSFTSVSTNSAIHLICIKKIIRMVIIQAEDYPHDDFFFF >TbOr28 MSERSEKIIDNEIIDRFKFYFWTLYPIGLWPKNNKSHLTDFTESVKIIFVLTVLLYRAVPTS LTIILEIKDPALAFKMSSAVIFIWTSIWQYSLLVRKSQEIYRCLENIQSFWKITQKIDEKESI LKFNEIGLRQAMWCAFSMHFAGMYYYMTVPYSKSIEAQHLIKLVVFYVVQFVSTLVVH SISVGVCALFALLVTNASAQVDIVMKRINNFILNFHADRKTNDIKYSKIFYQHWQTIRFA AKLETILSLVLFIQVCSCTVYICFLGFFFLTEAEDSQINNTISFSMILGAFISHMYLICYLGE GAETQYKNVGNAYWCTNWYNMPHQLQICGTMFILVSYIPLKVRAFIFKMSLSCFGDVSI IFSIVIN >TbOr29 MTDTSPETIDREFDKRCKFYCLILNPLGLWPRNSKSRLSDCRDFLKVIIVITVTLYIAIPSSL KIILEVKDPILILKMGADVGFIWLCTCQYILLVRNLQVISRCIENIRSFWRYTHKIEEKQSIL KYHRFGLRHLMWCAFIMHLAGLWYYIMVPYSQSFETRSLSQRLLSYLIQGISSLVLQNV SVSVCALISLLLVNASAQVDVVMKRINCFILNFQSDQTTNEIEYSKLFYQHWQTLSRFAA QIKNLLNNILFIQVCSCTMFICLVGFFFLTEMEDTQIVNTVTYCMIFCAFISHIYTTCNIGEE

427 FECQYKNVGNAFWGSSWYNMSRQYVLCGMLFIVVSSISLNLKAVIFKLTLSCFADVMR SSMYYLNFLRKTM >TbOr30 MNTKSTTKIFDEEFNSKCKLFFWILYPLGLWPNENKSHLTDFTDTVKKIFIVIVSLYVVVP TSVKILLENKDPVLLLKVGAAVCFIWMCILQYILLLRKSQDIYRCIDDARFVWKYTHKIE ERRSILKYHRIGLHQLMWCAFIMHLAGMWYYMMVPYARSFRTQNSLTLLIFYVVQCISS LVVQNISVGVCALVGLLLTNASAQIDIVMKRIDYFVSNFQPDRKSNDEIYSKIFNQHLQT LRFAARIKSILNVILFIQVCSCTLFICLVGFFFLTEMENTQIINTVTYYMLLCAFISHIYMTC YIGEEFEFQYKIVGTAFWCSEWYNMSRQYVLCGMMFIIVSSVPLKLKALIFKLTLSCFAD VLRSSLYYLNFLRKTM >TbOr31 MCNKRKSKTINAQFNEKCKLHFYILYPLGLWPSEDKSHLTNFTNTAKMIGIFTVLLYVTV PISFKIILEKDPELIFKMASGVSFVWMCIWQYSLLVRRSQKIYRCIDNVRSVWENTNKTEE KQSILNFHKIGLYYLIWCAFVMHLAGMWYYIMVPYSRSFDTQYTSKLVFYLIQCISSLVV HNISVAVCALIALLLTNASAQVNIVMKKINYFVSNFQDGRKSNESEYSKIFYRHWQTIKF ATNIRTILSLVFMQVCSCTLYLCLVGFFFLTEVQESQFINTASYCIVLCAFISHMYMYCWL GEDSESQVSIYKEIQNFLYYS >TbOr32 MSKTVKSKIENTTFDHKFKFYCYILQPMGLWPTKTKTENTFKMVLSVSLLLFLIVPNTLR IIIEVKDAIIKLKMFAALSFIWMSISQYVILILKSKDIYRSIDDVRIIWDNTSNSKKNLLMAR FYRNGLRQSVRSAFLMYLGGFFYYTVMPFSSGKTALNNNGTLSRDFVFSTYTKIFDPQD KITFPIVYAVQWLGGYVVYTISIGSCALAALLLTNVSAHINIVMHRIDNLILHFSDSANNN VEFSKIIQQHRRTLSFAARIEHILKYVSFIEVCGCTMHICLVGYFFLTEMERSQPISTFTYSI LCCSFISHIYLYCSIGEEFEAQCRKVANSFWMTEWYVMPRSYIFSGKMLIMASNNPFKLK AVIFSMSLGTFAEVMKSSLVYLNFLRQTM >TbOr33 MIDIFGKSLNIEFDQKCKLYYWLLNPLGLWPSRKKPNFVGWNTIIKIILSISLISFSTIPFLL RIIIEVKDPKIILPMFGLWSYMTMSMVQYIVLVRHLKIIYKVIDDIRNVWNITSNLKDKEA MNKLHENGLRHTLRSAFLMYLGGFVYFTAMPFSKGKVVSSEDGSVSRDFVCSTYRKIF DPQDALTFPIVFVLQWLSSYIVFTISSGSCALIALFVKNVSAQLEVVMNKIDHFVIHFDES DKNNKTNFAKIFHKHSHTCRFATEIENILYYVSFIEVCECTVHICFVGYFFLTEVERSQPIS TITYIMLACAFTSHIYLFCAVGDDFEDKYRKVGNSFWMTEWYLLPRKYLLSAMMLIRM SIMPCTIKAVIFKLSFVCFADVLRSSLIYLNFLRETM >TbOr34 MLFSIIPFILRIVIEVKNPMIKIQMLGFLSYLTLTMVQYIVLLGHLKIIYKVIDDIRYFWNYS SNSKEKEAMNKLHENGLRHTVRSAFMMYLSGFLYYTIMPFSGGKITDSDGSVTRDFVCS TYRKLFDPQDLVTFQIVFVIQWISSYFVFTISNGSCALAALFITSVSAQLQIVMHKIENYIL YFRKFHQTNEKELHLFAIKIQKILNYISFIEVCECTVHICFVVYFLLTEIERNQPISTITYIMI ACTYTSYIYVFCAYKEVGNSFWMAEWYLLPRKYLLSGMMLIRISLMPCTMKAVFFKLS LVCFGDVLKSAYMYLNVVRETL >TbOr35 MKTKMTDEYDDFEHKFKFYKLTLNPYGLWPESTKSILPKYGNKLKIVLSILLMLFNIIPN LLKIFLEVRDPIVKLNMLGAQSYYIMTVVQYVLLVSHLATVHRCINDIGTFWNQTVDEQ EQKSMLKFFKNGLHLSVINTIFVHISIILFFAILPFSDSALIKDFACPTYTKIFDPKDVLTFRI VFVIQFIAGWVTMNISSGFCCLVVFFITNLSTQFDTLMIRIIKFIRHVNKSHQVSSKGLSNIF RQHAQILRFAIKIEDLCNISAFIQLLHCTMHICGVGYFFLKGVDDSHVVGILTYVFLGITFT

428 CQVFLLCSIGDNLETQCGNVGIAFYMTEWYALPPKYRLSAIMMIRISNNASSFQMKAVF FNISLLCFVNVIRTSYIYLNFLRKTMK >TbOr36 MKTKMTDEYDDFEHKFKFYKLTLNPHGLWPESTKSAVMKCGNKFKIVLSLSLVLFNIIP NILKIIIEVKDPQIKMHMFGSLSFYAMTLVQYILLLWNLTTIYRLINHIKTFWNQTVNAEE KISMFKFYKNGLYHSVINAILLHMSVIFFFAVLPFTDSNMKGDFACPAYTKIFDPKDVLT FRIVFVIQFVAGYVTLNISSGFCCLAALFITNASAQLDIVMIKMRNFCVHLSGFELSNKSE LFDIFRQHSEVLRFAVQIKDIYNIAAFIEVCHCTIHICFLGYFFLSGVENSELLSTFTFTLLG AAFTCQIFLFCSLGDDLESQCKAIGSTFWITKWYALPRKYALSGIMVMKISNDTFSSQMK AGFFKLSLICFANVIRSSFIYLNFFRETMK >TbOr37 MTQSVKSNHDNNSDHKFKIHFWVLKPLGMWPSEKNCNVSYRNKFIMTLSVLLMLFVLI PETFKMIFEVEDPMIKLYMFGSVTCFSLIMMQYVLMVGNVTLVYRCIDEIRTTWNRTMD KKEKQSMLKFYKNSIKHVVMIIFLMYLSAIIFLGISPFATKFVADRDFAYPAYTKLLNPQ DYLTFRIVYVVQLFAGYILFNSSCGFCCLAQFFLSNISAQLDIVMIKIDNLILHFSESHLISE IEFFNIFRRHAQTIRFAVQIRELLNMGSFMEVCECTLNTCIVGFLFLMEMDNSNLIGTIMYI LFTFALINKIFFFCSLGDELNYQYKSVAMAFWMTKWYLLPKKFALSAMMLLRISNSSFQ MKAGFFELSFTCFQNVRISEFIVVTCTLISYLKNHFSGYTNVVDLFQFRQRNLKLNKNAT KSLWQIAINNKYIYYKIRRYFLPNRIL >TbOr38 MIHIISKNYDNDFDHTFKIYYLILNPLGLWPDKTKVILISYRNKIKIILSILLLLFVIIPHSLKI IIEVKDPKTKLNMFGCLSFFILSLIQYVFLLLNLTTVYRCIDDIRTIWKKTVNEKEQKSML KYYKDGLQHTVRSTVPMYLSAFVFLTIFPFSSKYVGNESSDREFACPTYTKLIDPQNLTT FQIVFVVQLFAGYVAFNASSGFCSLAALFFTNISAQLDIVMIRIDKFVLRFSKSCLFDKLEF FNIFRQHSQILRFTDQIENLLNIGSFIEIFQCTLNICLVGYFFLTEIQDSQLITTLMYTFLTSG FVSQIFLFCLVGDDLAHQCKSVGIAFWMTKWYVLPSKYAMLAMMLMRISNASVQIKA GFFELSRTCLGNVLKSSLIYLNFLRETMH >TbOr39 MVQSVIQKYDYDFDHKFKFYYWILNPLGLWPSKKKMNVVSCQNKLKIMLSISLLLFTAI PLMLKIIIEVKDPKVKLNMFASLSFYFMSMVQYLLLLRNLRTLNRCIDDIRNTWNKTVN VKEQQSMLKFYKDGLFQTVRSIFLMYLSAIMFLGIFPFSAKFVSDNALSRDFACPTYTKL FDPQDLTTFQIVFVVQFIAGYVAFNIPSGLCCLATLFLTNISAQLDVVMIRIDNFVLDFSKS QPKNYLDLFQIFRQHSEILRLAVKVEDILNIGSFIEVCESTIHICFVGYFFLTEIENSKLISTL MYTCLASAFVSLIYTFCSTGDDVECQCKSVATAFWMTEWYVLPPKYIISAMMLMRTSN NSFQIKAGFFELSLMCFGNVLRTSLIYLNFLRETML >TbOr40 MIHRVNKAYNNDFDQKFKFYSRILSLLGLWPSKENLNFGYRNKIKMMIVISLLMFNAIPL MLKIIIEVKDSKMKLNMFGTLSFFIMSMVQYVILLRNLKTLYRFIDDIRTSWNRTVGEKA QQSMLKFYKDGFQQTVRSVFLMYLSTIFFFGIFPFSTNFVANTLPRDFSCPTYTKIFDPQD LPTFCVVYVVQLIAGFVAFNIPCGFCCLAALFITNVSAQLDIVMHNIDNLTKTDKYSKKD LNEIFQQHSRILRFVVQVEDILNLIAFIEVCESTMHICFVGYFFLTEIENSQLISTLTYISLAI AFVSQIFSFCSIGDDLECQCKSVGMAFWMTEWYILPPKYTISAMMLMRISNSSFKIRAGF FELSLTCFSNVIRSSLIYLNFFRETMD >TbOr41 MHIFSDIHRILKSLKFLATLNLVWPLRFGQNEKIKSFIYRLYKYIYVSILLSTWIPGVSSFY TLRNDFSQLVKLLAEFIVLTELMINGLIINYHGGKMRELIFELEDFVKNTNFQEMMVVQK

429 YVNYHLSFFRVIFTSFIFAGITFVSSPLFMDDQQFPTDIRYTFLVASFPMKLLIYITQIATIFF IVTGFSCDMTVAYILWGTTVRLELLANRFLNAQTACDVNKCIKQHQHLLLFHEKINQSIT YTVLKSYLMMSIVIIAGGYQLINHEPFSIIARFILIVSASILRLYVNAKPGDDLKNISDEIKQ SIFESKWIEMPYTTKLSLLICMERLEKPMVLSVSGIMTELSLKHYAMYLSTTMKLFANIRI IFDREH >TbOr42 MYNNLKAAEAFRVVKKFLILTSIWPHNKKKLGRYKSLFYEVVWHLSLFNLIFLFTPMYL NIYFARDQLIEMMESLSVGETAGNAALNMILTKYFRDELQKALSEIRDFIHHPSEQDGKV IQKYVDKHKLLYILFCIVYFITPAIFVISPLVKHTRFPAAIWYPISSNSIKPIIVIYLSQTIGAY QIGFTCYVDIMVVILLSYTTMKFDLLNQELESVQDIGHLQNCIKKHQNLILFTNRVFRITR YLILDANILMALTAIFGGLLLLSHVPLLVVLQFVFMSITAYFKLYVFAVPADALMQMSEQ VTLSAYSSNWHRDAQQFYKPLFIMVERSRKVLVLSVPGLISTISLEQFGDSVSFIVSCIVR LRPFF >TbOr43 MTFSLSTKHAFNLVKFCGEITCTWPTPRSSQLRVMIGTIRWWLLIGNVLALLLSLAFAVY IFRANLTIATQAISEFGAAFEVLCNLFFCRTHTIRLQKALYEIESFIGVATYEEQMTIKKYT MRHKYLSAFICLSYVATAVMFISPFFTDNKLPADGWYPFPIDTFVIKSLLYVSQSIVILQS GLSISVDFMIASLLWYCTVKLELLAKELTRINDDECLKNCIKKHQHLTLYAASVNKIVRF MILKANATMGIAVVFGGFLLIRHAPMMVTVRFITFTLAGCLKLFVTARPSDDLKTTVRS LSNQNFIYISLHSGLFIFISQSQNISMAVYFSNWVDNSKQVKQNMLIMMKRSYKPFLIYVS GFIREISLLQYADVCVLITLSKHND >TbOr44 MTFIIWNAKQAFSILKFSCKIMCTWPRPRNTFKQVLFDECTWWFNLINVTALALPLSLAI YKFSNNLSVAFQALAEFCASMETVCSLILYKMYATRWQMSLLEIEPFLDKASHEEQNVI TKYANRHQYLYSFICISFIITAVLFTFVPLFTNNELPADAWYPFPIDAPLVHLLLYILQSVV IFQVGIFIFIDFLIAMLLWFSTTKLELLGYELEQIRNSEDISICIKKHQHLTLYAANANKISS WMILKSNAIVTIVIISGGFCLIRHAPIMITMKFVALLISGCLKLFVTAHPADDLKTMSQDIS MAIYNSDWIDISSVEVRSKMLILMRRSQKPFVINVHGVIKEISLAQYADIFPWQCYCGSV QLNLSYWHMSSSE >TbOr45 MTFMWNAKQAFSILKFSCKIMCTWPRPRNTFKQVLFDECTWWFNVINIIALALPLSFGIY KFSNNLSVAFQALTEFCASMETVCSLILYKRNATRWQMSLLEIEPFLKQASNKEQKVITK YTNRHQYLYTFTCISFITTATLFTSIPLFTNHGLPADAWYPFQIDAPFIHVLLYILQSIVIFE VGMFIFIDFSMAMLLWFSTTKLELLAYELERIRNSEDISICIKKHQHLTLYAANVNKISSW MILKTNAMITVGIISGGFCLIRHVPMIITMKFVALLISGCLKLFVTAHPADDMKEMSQDIS MAIYSSNWIDISSVEVRSKMLVLMRRSQKPFVIKIPGFIKEISLAQFAEVCFKLLDFRFL >TbOr46 MLSALLGFRIIRIGCTLCCTWPWRRSSRKMVFLDQLFWWISEINNLSLVIASFYYGYKTIP YNMSKGLQSVCEGIGFMEGSMTLLLCKVYSTSFQRAMAEIEPFLKNATEKEQKVISKYV NRHPYALTFLVISLFVCTVIFIATPIFTGDPLPLELEYSFPIDQLIVYLTMYLLQTMCLVQV SLSGCLDIAIATLLWYNITKLELLGFELENMKNNEDLKSCIIKHQELDLYARNIKKVSSWI IVKSIGVLGAVLISSGICVIRHVSEDVIVKFVMVIICGIYKMFVTSRPSDDLKIMSQQMSFS AYSTDWPNIHSSKFRNAMLLLMRRSHRPFLIGVVGLIEEISLAQYARFLTTSLSYFMRFK AIFE >TbOr47

430 MVSALKGLRIIKIGCIVSCTWSSLSNSWITIFLEQLYWWISEINHIMIVVSLFYSTYLTISYD ISESMQSICLLFGITEGIISLLLCKVNSSSFRRAFLEIEPFLKNANKQEAKVITKYVNRHPYL FTCFIVCILGCAFLFCVIPIFTQDLLPFNIKYPFLLDQWTIYLLVYVHQFISIIQVATSGCLEI ALATLLWYNVTKLELLSSELETMRNSEELKNCIKKHQDLDLYARNLKKTISWIILKSIW MLGIICITAGICVIRFASGDKIKFGYVIIAGAYKMFITSHPSDDLKTMSQQVSMSAYNAD WPSIKSSTFRSGMLFLMKRSHQPFLIGVVGVIREVSLAQYARFLATSLNYFVKMKALFE >TbOr48 MLTALKGFKVIRIGCAVSCTWPWLRNSRILIFLDQLYWWFSEINHTTIVVCLFFTAYASIN YDISESIQSICLAFGIIEGAITIFLCKIHSSSFRRALSEIEPFLKNATKREARVITKYVNRHPY TFAFFMSCMWGSTVYISALPIFTNELYPMNFKFPFFVDQWAVYLTVYALQFISIVQVSSS GCLDLALATLLWYNVTKLELLGFELENTKNNEDLTNCIKKHQNLDYARNVRKVTSWII VKSIWILGVICITGGICVIRVDSGGFTSENNFKFALIMISGAYKMFITSHPSDDLKTMSQHI SVSAYNTDWPSITSLKFRRAMLFLMERSHQPFLIGIVGVIDEISLVQYARFLITSLNYFVK MKALLD >TbOr49 MKNKVITATKFICATVAVWGCPMPMNCSKTGVFLHIIGQVTSVVSTALLFVAVGLEIYA SRKDMSVLPELVVFSTALHLIIVKLLILLVFQRDFQMLYHEAECFIKSATNKELTVLQYY VDKCKHFYFMMSCMPYMTVTLACVMPRLMPGKHLVADAVYPFAIDRNPAYELVYVH QIITFVQTAPSMCINNFTTFLLWTAVARLDILARQFQELTTTAEFRNIIRTHKTILRFIEKTI RGTRYFLYEDLIVVTISNIFAALLLYTSQPIAIKAQFIIITVCTITIIISICWTGDNVIDINENLG LALYHSYWTNMPAVTTKNVKFFGLKMIYPNYISAKPFIRILSLEFFGNVVSMIFKYFTAM NAAIMKD >TbOr50 MKLTPLSAAKITTTTVALCGCNAVPIRSKTNFHLYSFFKLLTFSSITCVLVPLIFRVYFSRN DMTVFPKTVVYSCGFVYFLLKFIVFFIHQKRIQLVYHEMEWFINRAKGKSLLFLQFYVD KYRHFYSILTCSPFVCSMSVSFLPIILREKHFPLETVYPFATDETSVFLLVYAQHSVSLFQL ASCMCTNNFATFLLWFAISRLDLLSTQFPHFKTPADFRKFTRTHQSILRFLDKANDVISYL LFVDLLMVTTACIFSALYILTKKDSVLFCQFNFCAEKKTKHIVLPHVVYFSCVFWLGSKQ SNALDYSLYQSDWINMPTGLVKNILFAKMEMQIPNVISVKPLVSALSLEFFGHVMAMLF KYCATMNTMLLQN >TbOr51 MMKLTPLSAAKITTAIVVLCGCKPVPINSSRVTYHVHSLIKILSIISVSCFLVPLVVAGYAS RNDMTVFPKTVVLTCGCVYFILKVTVYLIHQKRFQFVYHEMEWFIETANDESLSFLQLY VDKCRHLYLVTSCSPFMINMFLCLLPIFMEGKHFPANAVYPFATDKTPVFELIYAHETIS MFQVSSCMCSNNFATFLLWYAVSRLDLLSRQLEKFETAADFRKSIRTHQKILRFLGKAN EGIKYLIFVDLLVVNTACIFAALYILTPQPRVIKLQFTIIVFILITMLAMICWTGDNMIEISN ALEYSLYQSEWIKMPIQLTKNIVFAGLKMQVPNVISVKPFVLALSLEFFGYAVAMLFKY FLTMNSLLLQN >TbOr52 MSRIKLTPLSAAKFTIATIALCGCYPVPYQSSRKVFNLFRLVKILSIISTSCFLVPLVFGLYA SRNDMTVFPKTVVQTCGLVYVILKLAIFLIHQRRFQILYHDVDYYIKIAEDKSLAVLQYY VDKCWFFYMTMSCSPYMTCLFVCLLPYFMEGRHFPGEAHYPFATDRSPVYELVYLHQS FSAFQLASIMCTNNFITFLLWYSVARLDLLSTKLKNFLTNEDFGKSIRTHQIILGFIKSAKN GIKHLVFVDLLLVTTSSIFAALFIFTPQLLIVKLQFVIVFIASISLIVTSCWTADNLIQMSNK LEYSLYYSDWVNMPLASMKNIMFTGLKMQKPNVIPVRPFVGELSLEYLGYVLAMLFKY CAAMNTLLLQN

431 >TbOr53 MKVTPLSAAEFIVKTIALSTFYPGPMSSSRKVYHFMTFIQILSIISTSSFMISLFFGVYVSRN NIILFTKSVVEFCGMVYCLWKMIVLFFHRRCFQMLYHDVVGFIKTADNGTLAVLQSYV NKYQYFYMTMSCMPYMICLFISILPCFLEGRSFPGETHYPFDTDRSPVYELIYLHQTFSMI QLASIMCTNNFTTFLLWYFIVRLHLLAMDLENLLTAEDLYKSIKTHKIILEFIKTAKGGIK YLVFVDLMIATVSTIFSILYISTPQPLIMKTQFVIVFFSSFMLFFMPCWTADNLIEISNILEN SLYYSDWLNLPADSIKNIMFTGLRMQKPIVIAVKPFIPETSLEYLRFVLSVLYKYYTVMN SILLQN >TbOr54 MKVTPLSAAKFIVKTITLSSFYPVPMSSSRKVYYFMTFIQILSMISTNCFLIPMCFGLYVSR NDMTIFPKSVVQFCGLVYSLLKMIVLLFHRRRFQILYHDVVCFIKTAGSGTLAVLQLYV KKYQYFYMTMSCVPYLICLFVCLLPCFVEGRHFPAEGLYPFDSNKSPAYELVYMHQLFS TMQLASIMCTNNFTTFVLWYSIARIHLLSMDLQNVLTAEDLYKSIKTHQIILE >TbOr55 MEWFINNANDQNLAYLQSYVDKYRYFYSILSCSPFVLAILACLLPILMEDKHFPMETVY PFETDDTPIFMLVYAHQTISLFQLSSNMCTNNFATFLLWFVISRLDLLATQFLNCKTASDF RKFTRTHQNILRFLYKANDGIKYLVFVDLFIVTTGCIFTALFILTPQPSIIKVQFTFIVIVLIT MIGVICWVGDNMIHI >TbOr56 VTPLSAAKFIIKTITFSTFYPVPLSSSRKVYHFMTFIQILSIISTSCFMIPLCFGLYVSRNNM MLFPKSVVQTCGLIYSLLKMIVFFFHRRRFQMLYYDVLRFIKTADNETMAVLQFYVDKY RYYYIVMSCVPYMICLFICFLPCFMEGRHFPAEGLYPFDINKSPAYELVYMHQLFSSMQL ASIMCTNNFTTFVLWYSIARIHLLSMD >TbOr57 MGNEEFEKLIRPVRMCNQIMGVWPSNKKYGFIIESVKTFQLHFMFLVAIFHVTTTVIDM VRNRDNVNEATECALVASAGCLCFFRFFAFLSHRKDVAVVVEEMKSDWSLLNSESKQI LREKTYYVYRLAKIFIISILCSVVAFIAGPLLQSEDKILPFRGYYFINHPSPVQYGTMYIFN AIIGFFTTNTISSATSLSLIIVMHGATKFSILQMKIKSLQVASYKTENKMNNLIREHQNAIR FAETVERILSLITFGQFIISSILLCFAGFQMTMLEDKRSQIKYILFINSAIFEMFIFSYSSNQLI TESEKNCYSSYETCWMVMNKQLCKKIMFLMMRSKKFCKITAGGFVDVSLKTFAGVLQ TSFSYFTMLRAVNDD >TbOr58 MSSKEFDRLIRPVRICSQIMGVWPPNQNSNFIIKWIKYYQLNFMFLVAIFHVATTTWDAV RNRHDINELTDPALVASTGYLSFFRFIAFLTHRKDVAFVLEVMKSDWLSNDNKQLLRDK TSHVYKAAKIFIIILQLSVFSYVLGPLLKPGNHDILPYRGYYFVEHPSPSQYKIMFICNTVI AFLTCNSVSTATPLSLVIVMHAATKFSILQVKIKSLQDVNSDTAGNLNNLIREHQQAIRF AEAVARIISLISLGQFVLSSVLLCVAGFQMTMLHDTHGFIKYGMYINSAVLEMFMFSWSS NQLMEESEMICLSSYESCWIIMNKHLCTKITVLMMRSKKFCKITAGGFVDVSLKTFAGV LQTSFSYFTMLRAVNDD >TbOr59 MSKKKLNEGNDFLNHAGINAKLLKMCGIIKLESEVLTNTLALFSFSCLTFYSVTYTLIFV QFINNIEAATAYLSLVMSFLGGQSRFSLLLWSRTRIMTMLDKCDSLWSTLTAAEKQTIHF FLRKSAIITYIYLNFCAFTILSYAVAFIFQHQAIDENNQTLARKLPYAFLGQVTRTPYYEM TYAMQMISVVNVGFTCVACDTTGMFLIMIACGHLSVIQQRFFQFRPPVTRNNYECLRKN FNRQEQIQIISTNIVYHQSILEYCKDIEKLANITFFTGLIGTTYNLSIMGFKLTSDDPAKMTF

432 AFQTAILVMQLFVYNWPPDLLINHSMQISRAVWYSPWHKLHTESRKLLTFVIRRANRPV QLTAGKLFYLSLENFASVKQTTEQTIKKKRTSTFALYNF >TbOr60 MGFFKRKFKWRLANMVEPIMKSTGNWYYEDNSLGGQRTKFALQSYNFISLFASLVDNI TDCYCVWNTNKGRFLDNIFTILVIIVALVKLIVFSILREDVKYMLKFGINRFWKRTYDSD AEKLIISSEKRGFYMLLFYSFGVLFSTFLYVTRPLIENAMIFESERVRQLPFPLHYGEETFV TPLFEFCYICEVIVSCQVCQLTLTFDYCVVAFIVHSDVQYTLLGRKMESTTETIETLISERH DCNAQKLLNDLFFTLRNHVRCHNELINFVEVLKRIFSVILLIEIVLPSIVICVGGYRILTYPN SFGMLLCIFTTTTAVLEITTFIYICDNIAVSSKNMAFSVYKILWYELPICRLGRRIRYVIQM SIMRSQKPCYFTAGQFDVVTMKTLTGVMKTALSYLTLLRTLNE >TbOr61 MRKYEYTTVNFCHFFGRFVGIWYDETKFQRIFEKGFLCYATYVSMFSLLVTLYDTWKS RRSHLIFHRISNNILAIIITIKLMTFIYYRRVLKMLIRFSELYFWDCKYNDVQNKIIRKFNKL GEKMLIYFIISTMSCVIAYCIRPIVDNLSRPKGERLKRELPFNLYTPFNLLASPLYECVWV MAMVFTSNYGLITFSVDIFLSTFMVHLAAQFKILGQQATEIVTDFCMKNADDKFQEYLK ILCSELKLFIRKHNNLILYTKRIESIFSSILLTILVLAGIGACFCGYIIMTEPLLGIVIVSFVTA SSGLVQIYIFSYICTTIVDQSSEVCFKTYCSLWHKVPSEGYGCKFKDDLIFIMMRSQKPCV LTAGKFYAVSLNTFASVVKSSVSYLTVLRRARDK >TbOr62 MKTDIYNLGTFKLLRVFMKIIGLWPTNNSKEQIISNVCRTNYVFTMVVACCVEFPDLYFS ARDSLFALAVSAPSFCGVYSESLKLFTFIRRKREVLEFSRYTENEYWRANYNPEETRIIRQ CDRLTLPSCIVYVMLVQIVSFAYIFLPLYESIGQHDSNRSLPFGCWFEYCRVSPIYEIVYFL QSSGCISVSTCFSSFAVFIYSMNSFAAAQFRILQQNLKKIFDSTEDKTFYQLTKTLNYDND YKKVIVLKKLIKKHQELIAYVKKIDELINVIIFFVVATCVVQIVFLWFQITLSENMTILRKL LATCHLIGLFSQLLFYCWSSHQIILSSIEIRESIYTSKWYLMNFYIKGKEISQCVRLIINRAT KPCILNCGVENFNIIFHIFKACWLNRKLIIFVIYCGQLLGHYSKNIEWLTFYFYKFLIL >TbOr63 MNKTTSSIENHKCQKMTKFFMRLLGMWPSDTPGEKIVADSVLLLAINMMVISSIAMTFD FWFQCNNFYEIVYVIPCVSVTYAELAKLLVFFFNRKNVLDFIMFTQNYFLRRSYKQEHL QILQNCNNTNYPRCAILICIVQGVTWHYIFIPIFENIGRDESERTLVFRWYFDLIPYTTSPY YEWVMLVQVISTFYVGVMFTASAIFLFTINVYSAAYFDILKNRFETLCEFDELTDKKMN FTETEHLYYKKLIACIKEHELLIQYVNKIDYLYRSIMFVQVFSSVMQICFSGFQLLLGVST SIQRTILSGEYVIGSFVELYFFTVSSDEIKFSSIEIQDSAYNSKWHSLAFNKFGKNFRTSLAF VIFRANYPSVVHMGSFGVMSHQIFTWVLRTSGSYFTVLRQFMEQSEINV >TbOr64 MSSPAREKAKRYLSVSLCKNFLSFVGLWPHKSEHVNNLVTFMYLHTTFALVLSISIVLK GLSYCCVNIIHFIRLFSVMSSIFFINAKFVCMSLKKNDLFRIIEFTLNKVINISYNPVETKIM VACDKYCIKFCIVLLLGNQLTVLSYVGYCIDEWARGNKTLYPFPYDYIFNTPSFFRTTPA YQMLVVHQFVVNYIVGNVYICFDIALVFFITFARGQFEMLHHRVSNIYNKSSGYVDITTK LKSCIKEHKISICYVKNVNNLFSIVILMQVLFLSIIICLTVMEGLMIEETANTNIFLRISNVS YTVVAFVQLLSMSIIADNILCSSTNLRNYLFHSPWYTVHLKEANRVKSSVLTIIMRTSIPC TLTVGAFSVVTVETFTKVISTAYSYVTLLQRQVDR >TbOr65 METIQNQNCVLLTFVKYYLLFFGFWPQKKRSFFNFTRFRLHYSAFALFCTNNDVYSAIL DCCTNVLHFVRKFNVVTSILIVDGKFISIYLRKTRIYQLIEFVCDDKRGSFVSEKTSKFMT PHDKYSLKIIIVLILGNQITIISYISNLVYTWFIFNKSERSFPYNYLYSHYPEFTKTYPLYEVI

433 VMIQVFNNYIFGNLYICFDICLIFLILFVQNKFETLRIRISNVVAIPFDCDNHLSTTKSKIRE CIVEHQKLIKFVQVIEYSYSTVILMQIVFLSMILCMTAFENLKAEDNVHLNTFLIISNIFYT LTALMQLLFTSNAAGGIMTRSTNLSIATFHVPWYSACTRKQNDITSSILFIIMRAQKACKL TAAGFCVISAETFRMVLSTAFSYFTLLETQT >TbOr66 MLRLKIKVNNRFLFNFIKYYLSFFGLWPYKNNYCMTIFVQICLVISISALVLTNCNVIRIFS FCCFNVAMDCNLTCSKIMDDCNKRTLKSIITLFICNQIVIISYVAYCVDEWLSIGRYSRLF PYEYFFKLPNFMTKFPLRNIMIINQFFMNYIFGNLYVCSDISLIFLLTFTQGQFKILRERISEI GIKPNDSFKLLKNMNDKLRACINHHTILISFVRNIEKIFSVIILIQVVFLSIILCITVFESLKVI ACPCERNKMITLIICMVLQLLIMTVTADDIMSMSSSLSTTIFHVPWYFLDSDVIKRMISFVI MRAQKSCTLSAGGFCIITVETFKMVLSTAFSYFTLLQTKADK >TbOr67 MANRGKRYISICMIKALLIFTGYWYPKTELQKDLYEYILIYVITATVILPVITEFFELYASY DDPYTLAYTFTGTTTLYLIVVKVSVFSWRRKKILDLLKFTDENYWNPANDEFSFNLFRKI DKRVTIMISTFSFFASGTVMCYILAPIVENIGKKGSSRVLVFNGIRSFIPVPSPFFEMLYYIQ IIALFHLGATFMCFDHLLCILTSHVYAQFRILQHRIEMLGQKFIVISSTVGISSAKVSMSVK DFYAELKKCIRKHNEIIWYVSELESHYSFIIMSQIVVSSLMLCVVALEAIEMELNFVRRLT FINHMFGDFVQLLFITISCHQIMTESAAMSHAAYNCYWNILPFDSTGSTIRRTISFIIVRGQ KPCSISAGGFADVSLNSFTSAVSTSFSYFTLLREFL >TbOr68 MSKFTEKYLSLRVIKTLMIIAGFWVSKTKKFSYFLQFCFVSTSFNLLISLWIGVQGTTSSII QKDIGAVAYSTCIMTPVLLGVMKQIIFKVKEKKIIHLIQYSQDNYWYNETNLFNVNILKD VNRRSIFLMIFFTCSAMSTVTCFTVSSALDNIGVSDSEKILTYPEFETLMDLIFQMQFRKYI EKNYVFFMAFEYFVSITVALSFCIFDNMLLILTKHSAAQLIILGHELEQTLDKYKYDDDLE DLKDIKLRDLIKRHQDIIKFVDQIEDLFSIILLVQTVVSSFMLCVVAIQAFLMEYPFTKRIIF VFHMCGSMSQLFAVTTSCHELIHESYSLSMTLFLSRWYLHPFRKKLSYMINFMIYRAQK PLRISAAGFFNISFDTFGSVLNTTASYFALLRQFL >TbOr69 MTARKKYLSNKLIQILMSIVGFWIPHSNSYRYLLNFNRGFVLFNATVSVYMACLNFAISL GNPKTLAYSGCLALPVLLIMIRLIVFSMKKKKVLDLIQICDNCWRNENNPNNVQILEQID KESSFIIIVFTISCIGTAHCFLFAALIENYEKEKSEWIFIFDGFDLMIDRNFLLNHFTACMIC QYFATVCIGVCFCVFDNALLIFTKIARAQLKILGRDMNELNKKYLESTKLSMKDLDLKSS IDKHRMIIKFVKQVEDIFSFILFAQIIISSVMICVVMVQLFLIEYTLIKRIIFVFHFLGSFIQLY SITQSCNQLVLECLRLSDTIFFTIWFLLPVRNGVRYATKFMIMRSQKPCFISAAGFIYVSLS VFTSVNKTFSDKKKCIHVLNLLLLIFSIHVITHI >TbOr70 MMTTKQNYPSLSLLKILMSSVGLWITNSRSYRLLLQLNLYFAMLGTFIGVIVAISNFLVS LENAQSLAYSGCIAAPVIVVTIRVTVFAIKKKKMLRLIQFCHDNFWYNEKNIYYKQTLRE VDKSSILIMILFTMNTLSASTCFMIVGLVENYGKDKLEKKPIFEGLELYFGMDHVKKYA MFYFSFQYFGAVYIAVCYCVYDHALLNFAKHAAAQLRLLGSEMNRSDKKYFSSDQIFPS DVREKNNQMQSSIKQHNKIIQFVKGVEEIFSIILFTQILIASVTICVSAIQLFLVKQYFLKK MIFVFHTFGSIFQLYVTTSSCNELKEESLNFTTMIYSTAWYNFPMGNSIRYAVHFMIFRSQ KPCYINAAGFVNVSLDVFNTVINKTISIYTVLRKCM >TbOr71 MAKIKYHSVIITKIFMQLIGFWPFENARGRKISDFALINMFVALSVIAVIESLEFSHCCTDF NEAARLGVNVTTYVIACFKVVIMTIHRKRLLNLIRNIQKYFWEINYNNQDTEILRKCEKS

434 AMFLICVLSFLNQLTMWSYFSSFVVENIGKNESLRELPFKIWLDIPTSRSPFYEIFFVIQVL TVYFISIAYASFDLCLCLMCIYIVGHFKILRKRLVNFNKYNNKSIDSLSILKTHIQHHQSLI LYVDHLEVVFNLILLLQVTVLSVSICLVGYMAIIISDPVFLRLTFVFYYVAALFQLYSFTM SSDGLTRESCKVSSDVFHSNWYTLNENNPSLKIGRYIQMIIRRSTKASSLTAGKFFIVSLE TFTMVLHTSLSFFTLMIQTNFDISSEIGRASCRERV >TbOr72 MITVYLGFFGLCSVSLKNKRSILPIFIDIMSDLFGLILVAIKAINFQLRKTKFNYLLNESVIF MTVFTKYSDVEISEYAYEHCFKKVKMFFVLTGVISLYTIIQSFIIPIVERNHYIASNFENPL LDLSSPIAFYMIFVMQSITHSGMIFAIFGIDTFAILMLSRFELYSTFLVANFENSFYLTKKPS LSCTESEPAYSHDNDDEIKLHTFVIYERKKEIVNLETNFISLLKYNVKHHQKLITFVDILNT IFGSTLMSQLTLSSCMIILVTMQVTLNLGTPIQFTKYLLYFSLNLFHLFMLCNIGSNLIQTS DALTRAPWRSHWEFAELDSELRFLILLPILRAQRELKMKALHLAISRETFITLMSNSFSVI VLIRSFMASKIIA >TbOr73 MKLMPIPFGIFSICGFWRPVSWNSSLAARLYSIFTVFMYTAILTFTLSQMMDVILTVNNL DDFAGNSFMMLSMMNVCFKMTCVIRYRKAILNLSDTLTKEPFKPTDKYEMDVIQQQTR FARKLTFYNGSLTLATVTLLTLKSLFGAAKGILAFRAWIPYNYEKPLIFWFTFMHQTVAH AVGASIQVANESWMSGMMMQTIARITIVSHRFKKAAEVKNNASNDNRKSKIHSVESCIK YHSAVIKQVYQDFTEKLNKIIKTVVMMQFTMSSMIISLSVYLLTTLPLMSGQFFGLVLYF VAMCLQIFMFCWYGNEIILKSSSISNILDKMEWLNLEIRNRKLLLLVIRRSNKPMKMSSY FLVTLSLNSFTKANLDLTSQSTLINTLEFIILFHFFRS >TbOr74 MSTSTLLRNYNKYVRHTQLLLAIPGLYLTAKNRGKLTRILPNFISSLVIVSFFFASGAFTF KNYSNVALLIKDLSLTIWFLSLFVKFKCFTFSKQDLIVLNDELREAYENDLQNSVLRQIM LSDLPYFYYPNIINSIELSALVCSRVVAYMIFYITEGRHPKLYPTVHPWSAEPSTVLYEIDF VYESLASFTLLFIAIAVDVSFSLHVFKIYGQFQVLAYKLKNLDFGEKHDYELKECVKKHV QLINFCRCLARVYGEVVFFIYIAGSLSLCMAIYGGFETFKFRRNHKLSTAVSKYVWLLIY FVMKLEQIFLYAYYGYCITHKARKCQIVATFNDWADSEYKKIYKNLLIIILQKPVVLRVG YLEINIDMLKEAVHTALSYFFLLKTLDIE >TbOr75 MSTSTLLRNYNKYVRHTQLLLAIPGLYLTTKNRGKLKRILPNFISLFVMTSCCLAVGAFC FKNSSNVRVIMKVLSFVIGFWSILIKVLSFAFAKRDLIFLNDELRKAYENDLQNSVLQPV VLSHLPYFFYPILIITADTFMLWVFNVLTPFISLYKKGYHPKLYPTVYPWHSEPSSVLYEIE FVFELIMSWYSFIIATSIDASFGFHVFRISGQFRALAYEIRELSFGENHEYELKWCIKRHVQ TIQFCRHLNRVYGVLIFYLYIAGSISLCMIIYAEFQLIKLAKIYKWTNIIWKALWLVVYFS MKLEQLFIYAYYGFYITNESTHCQVSASFKDWASGDKEKYKNLRMIIFQKPIVLKVGNL EVTLKMFKEAVNTALSYFLILKTLAIE >TbOr76 MTSLKEYRKFSRFIHFMNSLVGYHCFLKTTGTTQKCLIFFSFFTSIISTITTINCCILNITRLS IFIKAMGVAGSCSTITIKLFCALYHRKRLQDLNSDLVSRFEKDLKQNENVPDLLCRMRQF SRPAYFYTGILSYIGTMYIILPIIILLKYGKYITILPSSYKPFNIYSDLRSYFAVFVIEFWTEIG MCAVTTSTDALFWMYLAIICGELRLISKKYDSLSKDMTNYQKNLRDCVSRHCNVIRHT KNVELIYGPIAIWLSVSSAVVICGQVLQVTQAPSILQLVVLFVYSMIKLVQVLLFSRGSEF VFTESENCLYKAFTSIRYEQTNVSMMRNMVVITAQKPLSLKAFGFFRISLELFSRVINASI SFFFLIQTFESK >TbOr77

435 LIVRLTQGQNVLIKSTKCSTSCSVFVFATEQVRPNCQHKKSPIHNMSSKSKLYDHNFNVI ESHFYKLNKCLLTLFGQWPWMNKFCKVLLSSILLTYLMIICYPGLRSMYLNRENIESMT RAFIPTTIIVFAYSKIFTYTIKNREIARLLESVRDDWYNLTSVDDKQILIKKLRIYYRVTQT YSISGIIGGLFFGMGPLIPKLLDIISPLDQPRPLRTVFNTDLSIYNMNIKINPVKTLSYGAINS LIGTTANCIFDSVYVLFVIHAASMFDVIKHRLLLVGNHLDERRNNFEIHDQQVHEELRFII SFYDRTIRYCKLIDDTCTWYQLVILGNTAFILAAGAMQILVMEKAEGFRFVILISGLVLHL FLYSFTGQMMLNRSHRILTDTYNAPWYKLHKTARQSINILQMRNLVPCTLTAGKLYAM SLDNFVAVFD >TbOr78 MKYEGKDVFDGVYFRMFKLTYMIAICWPYISIEILVLNLAIKICVALTMGFAQLKEILQID YLPDVFYACSQLFLNCIALILLLDSFIRRKQVKIFYEDIKSYWQIDKSGLHSMIRIKYAKM CKRVLVAYITIVQGSVVIYMLTPLLPIILDRIMPLDSPRKLEQMILADYSSFLPRFIYENYW IIFVNNVFQIFYCMQQVQVLNNNIFLLTSHACSKYKFCGESLKRIGLSSNRNFNFNKETIF FETRSCASYHQKALQQAQNTSNFFKISWGFGKAANLALLSLSLYGIVLLDIEQWHEIPKL ATFAYSVFFQLFIANFFGEMLRQHGRDILNSCYESLWYELPKNCQQNLLMIITRSNVDCP SSYISAYRKSLNDFVSSLNTSYCFFTTFLRMAH >TbOr79 MKQENKMPQKYADIKGNIFDSTYYKLYKYIYLSMGYWPWQSLPLSLLIYLPRLILSWSA LYPEFMALGKPKNLDEGIYIFSQMVIHGFSYVTIIDTVLRKKMLKRLMEEIQNDWDVAK SDESIETLHKYGYLSKWFMFIHMSSIQFTQLVYVLSPLIPKLLDRIAPLNESRPLLPIVYAD YTVYPDYISQSYTIIYIHNLFCVYVCAVQFQILNNTIFILTYHCCAKFAICGSKLQQVGVN GADGVLSPRSEKMVFAKLSESIRHHQKSVIFATSNSIYFSTAWITGKMVNMILISVSLYAII LNLHNFQEIPKHFSFASAMLFQLFICCYFGETLHQHSKRIRFQSYESIWYNRSKHFRKNV HFIILRCEQRVPPCSFTVFTKSLENFVSTVNTSYCCFTAFVSLSA >TbOr80 MAKHFLMFGYDKMRCMKLFLIVGNMYPFKSKLLSGLNFIICTSMMSILYYKMIFFWKH AKFDSLYSKVIFMVVNVLIILNMSTFILFQFYKNNIRRLFQCIENHWSNHEPKIHDKSSLH GTKMRCLRDELKFYIHLYLIYMFGFQMVNFSITMCRVILSIFMTSVELPTRLFYPVFVLNT KTKFDFCLTHLYQFLLFVIGLGASFSFEVLLAVFLAHNCILFTHAREKLNLLEKSMETVS KLKNNVKYFSRDYQESLDYFNLYQKTFSVATFARIFAILTITIALMMFAMISTTKSQLYR SISGISRTAVCYFFVFNYSEKISIYSNFSSDIYCISWYDLKPKSRVMINQLQSRCRRVCYVK ILGIHIVTNEFLTDILHAGMSFLTFCRRVLRF >TbOr81 MELDINQDYFIPYKSMFPKFKKKSVFKSIYKFLKIAFMVTGYWPGQNKKIGFIIRSSQLVL FQYIIYMLTYDLLELRYNVVNMCVNSIWFFLVFIMWIMNINNMNSLKHLKEFLSMIDIH WQLKALRYPHLQVNRQLIQKKLIHLLGFLILLVFIVLNFAPSLFLSFCGKIANTDASCSFQI RIPRIHKSPIIDILFQILLAPISLAYYYSFVIFAIAMLFTMIDAIYFIRYEMENIAYSKPNNNCI SIETSSLKTIKYSIKQHSELARSVKAFTAYVRLHLIFTVFIGILLMTFSLYALYHTSTSQND KVFLILSLTSEFLIVLVFGIMGELIIETNGIVQNAIYSSLWYEASEKVNKVGLIFLMNCSNV LNLNVCDALILSLDMVIKMSKTVFSYCTFFRNLENI >TbOr82 MFLNMLGPRFSWMKFYLGTTGLYPYYSVPFRALLYIVNMAVNMLMLWQLFILWHGM NFLTGFFYLSTPTCCIIASCAVYTYLHTQQTGIKQLCEHLKTYWGNIRKNLQNVQSLYM VRNRNKFLVWLLVYYTWIYVPITFQLAKPFKEKLKQSLTNSSEHVRLVFPITLLRTRTEY DLYLTLVFQTIFVYSACSIAVGYETLISILLDYNSMLFVLLRRKLDKVKNMTYNSHASKI VVDEYELKIEPTSLFQCIKLHQNYLNFFERAHDIFRPVWFSRVVVSSFLLAMYCYQSILF

436 MSDRVKLVRYLTSAGRVLFGIAFIVKYSEKLIKQSSMQDAVYSTLWYAHDKCVRQVVH VMQARAGKPCIISIYGICTISAEVFANIVSLGVSYFTFMASIA >TbOr83 MKIMQLKFFRVVKCLSCCLSIWPFQSKYVIRICGLTLWLYIISSFIFYQSVHLLYSSLSKEI LLYCLVLVLPSTTIIFLWATLNLNSTKIKMLLTEMEAFWNDVEQDEKYLKIFMKYLLRST NIAVFNAAFSISSIIVVLLGIGMRIKISFQMFKGTKYAIFLSNEVFFVSRIVFILYQVGFISVC VGIASLLVYSSAIMFTCITHSACRFYILGLKIESITEGYEKLDFKLIHRKLYNFIKDHKNIFK YQELLQSVLGKIFFYFMGVATLQMCLTIYHIHQNLRNHSNIGESFTKFLGTFVSLFLITLM CQQVISASIKFRDLVYFCEWYMFSKKTKNLYIILFMRTIQPSSVVAGKMFPMTMETFCGII QLSMSYFTFLTSIT >TbOr84 MKMTFFKKKEKSSQAFDNKFFSHNKLIVSLMGQWPYQSKLIRSTSRAMIFVYFLYIIVYD EIYGVLTIQMDFDVSTKYYCTLLADTVCAIKWVITLTNMSKIGRLITDLESFWSYVTSSR EKTLIRYAKFSRKLSLFYYVLWYGTIIFLYSIPSMTVVPIGSVNFDGNSSVVLEFGHEEIED KQTTYETLWIIVNVIVYPTLVLAEDTLVQVFLVHCSCRLAVLGKQITDSTTNEVSEMKSG VYKKLMIYLKDHEQTLNYVGMLQYVYGLFLFCDIGCHVGMMSIQFYQIQIHFQVNNTS FIGDCSLITLHILIMYFFMKPCQMLIDHNLRLCDMIYSSEWHSVTQKMKRIYCMILMATQ RDISIIAGDRKSVV >TbOr85 MKFLNKKLQYHGAKIFWKKRDNGIFDTRYYINYKRTLEFAGFWTNQNIIIKLCIWAVHL SASIAIIPTQAKYFLRHWREFENVAPNAIFLIGTLFLFVIFFINVLKSNELAKLHDTINLDW KTCRNQQERNIFQAVLIRGHKVMNLFRGLFVGSLILFCITPFTSQILDYIIPLNETRPHTHIF DIEFIYPRKYVFWRNYIYCIIGTIIGIFYYQVLFNEILVLVFHACAKFNLLQYNLEHIGKNH SGKSKSYGEYVTCDMKHCLKKHLDSVKFVRIFMKFTSLQSNTGVFCNTVGLSLLIYAIY LKLDDLSHMTKLIPLFVFTLIIIFQGCATGENLKSDSVEIRFAGYKSSWILWSTGNKRMLN IFQVVSEKPCAVPCIWVCDFSMEFFLSIVNTSYVYFNALLVSYKN >TbOr86 MKYLQNKSQYFLKNQFNKKKDNGIFDIHYTYYKRILEFSGFWTNQSLYIKLCIWACHVS AGILIVPCQVKYFIKHWKEFDNVASNAIFLAGTLFMIIIFFANVFKSDKVSKLHDAIDVEW KTCKNPMEKSILVEVLKRGKSIANLFRGLTGASVILFCITPFTSQILDYFIPLSEARPHTQIF DIEFLYPRKYVFWRNYIYCLIGIIIGQLYYQVMFNEIQVISFHACAIFNLLQYNLERIGKDH SPHSEFYDEYVMHDMKRCLKKHLDCIKFVKNFTDFIDFQSNIGVLLNVGCLSLLIYAIYL KLNQPSHMTKLVPLLFYCLIIIFQACANGETLKTDSAKIHDSCYNSAWLFWPTSVKRILNI FQVNSEKPLAVACFIICDFSMEFFLSIVNTSYVYFSALFVGKVN >TbOr87 PGWIPTVHNKLLQKSSKLGVGVSCSLCSVQVLIILAFCEIRLPQLYDCLRLVSLDKTSDPH VLKHYTFLFIAMFHTCMPNIIVYSTYETMLVITMFLRWNLIETDIRNLRSYDSEEIFAKEC KFYVLDHRQTVKLIRLDNDLYAALLLWGVTSFVIAMTVAIYNLEACFKNAEKAMVILPF FCITQVSIFCLCWFGQRIIEQNDRIKYSIYESKWYDLTKKQNVMMNIFQMSCFHPMETRA YGVLVLGTELYNQVMRLSFSYYTFLKARGEN >TbOr88 MSDDKEAPKFLRDFYAKDNKFVSIFDTHYKWMKYVYKGVGHWPYQNRVFRYVYYMT QMIFSIITFNGQIRSMMILYETDMDEFNINVLGIIIHATLMLFFFHNFCRKTHLEYLFDGLD KSCHYIKNEQEGMESILMFEKFMKIMMVIVGSMVVSFVTLPFALGIFKCNSNEFSDCIGK YFLIKLDYPFVVCYFLFLCAAFAGFCYINAFIFIAAAISIHYFVEMKTVKAKICTFGNLQKS ESLLFQDFETLVKKHNQIISAVQYMSEFFSILLFFGIGLIISGLTFTLFLMNKINGSPDRIILL

437 FAAAFVLLGVIGGCMIMGDEIKDGNDAIKIALYCSSWMTLSKSMTKMINIFQMHCNKPF RIHAYNIVELSADLYMSIIRASGTYYTLLRTTKN >TbOr89 MKKFRNTCSAFFEKVTDSAIKNKKSVYDNEYRWLKYINLFNGTWISQNKNLNLFIKYSH LLNSITLISGQCYYLSTLKNDIGDISRTGAWLAVSIYIALLRFNYIWNEKNIQNVLNTVNV QFQRIPNKMHGLCENMIKYSTAFATSIVSGEIYLPLRSYLKCRNVRKNLNACICNTVVIH THYPFYVIHWLIFVELILGVSMIHNFNCYAAVTSTEFFVHKSIIQYLEQIGTTYTELDDLEP ASENQLRLCIWEHKLLAAMKDANDCFRFILLYGMLINVAGISFTLFSIKHMIQTPPIDILMI AMFLPILTVSFIIVGVLCLVGDRVVETTDDFKLSYKSNWIVLPTKIKTMINIFQMRLNIPFV ISAYGIFDLSIQLFVQIVRISISYYTILLSL >TbOr90 MLNVACLFPDMNVKKNRSQSAPDAQLPERTKGSIIENEFRMLKKIFILLGVWPGQNKHV RMVLFVINVFVSINMLSALFIQVYQNRHLFDEVCSVSIWIVGFSTGFVLFLTFSTKTSKTA GIFFKADDWFLLKKSSIYHQTILDKLGRTGKLINLCTMSNIVVGLASAMVPLFLYFWNCK IQTTDDPQCMADQLVVKTNYPVAVSYVMFAIQTVHCTQPAICSQMIFCIFIITNSQSVSDI LQLHLPSIGKTCQSEEQNHDQLKICISDYKYLVNSIKLSNEMFAFALLVGITLVVIATTLA LYSLSEMSQNMQQSLTVMPILVIIFVIVYIICFCGDLLIQVNSQVKYAICIFRYRSGWMNL PKKITRNLNIFQMQCNRTLDLSAYGMITLSMDLYLQVVRTSLSYCAFLHTMKNHRI >TbOr91 MEVAIKDNPIFKEPPTRESSIFDTDYISVKYAYLYTGKWLSQKLYKKVLIHICYYIYTFFIF VGQIYYTYTIRTNFDELCINFIWLTCELLGIVMVTSEVFHHKNINRLLHVVDDHEKLLLQ GKYADFAKEEKDFCRKIIRIFEVLFFGIVNLLCTVPYVFIFLFCDLNSPNIAACKDQFIVLK THYPFGYYLVFSVPMGLYFVAYCFEFCCFVCIIAVQFFMERSIIIQELHSIGSEVNVSNLA QISKNKLRYCAAHHKILTKFITILVQCFSSVLMIGMAAEISCLTLSLYILSIVYANPERLMY ALPIMCICVVVILYLCRIGEKVRNTNDEFKNALYNSSWINVSTQLRSMMSVMQIECNSVS ELHAYGIFSLSMELYSTLARMSLSYYTALWTFRGETT >TbOr92 MHTKSSDKNHHKVDDFSLENSYYFKECKYFKYSLMATGVWPYMNKRIRNGLLTYQLI MLVLFQTGQWKHLIQHRSSIDEVCVNTLWIIVTSFCTVLVLHNIFYNQHVSFLLHMVAY QHEIRKTNSKSYFNTIQLFKLNKKISAIYLYVCVLSTIGPILFGFAKCDMKSSDRLQCMQK CFMVKSSLSFPTYIITMMSCYVFSYINVFNLCCLVGNILVTAFSMKSDIINYLNGMKNES DYENGIVNYDYGILKDNTHECLKQHQLIIRMLDAANACYAMPLLIGLSSDIITITLNLYT MNLFLHTPDKIVTIIPVITVSLFMTYVLCIAGEMIIEMNDILTHAFYSSGWIQFPKRILTTFQ IFQMRCNNRLCINAYGLIILSMEFYVNMIRMSCTYLTYLWSAKDESG >TbOr93 MINKLIDAKLEAHDSYLQNSYYFNQCKLIKYSMLITGAWPYIKKKVRIILYVSQIIAMCTL LGGQIEYVYQNRHNINELCFSTLWILNVACVTGHLFHNVFFNKHVFALLQAVVVVYEV EKSNKKSFLMMVPLSMIRKTIAAFYCSINILIIVGPILLGVIKCDMKSSDRFHCMKKYCIIK TSFSYKMFLIIIISACICGFLNIYNFYDSIANMMLVLTAARKRIVERISNMKYETDSKSNNE NSDVVSLKKQAQTCFKRHQDIIGILDVTNDCYWMPILICIGINILSIALTFYTILTFIDTPKL VLPLIPISILCCSFPYIACAAGQMVVELNDSFKYAFYNSQWMDFPKNIQTMFQILQIRCNN IIMISSCGIIVLTMEFYVNTMRASCSYFTFLWSVKHKSA >TbOr94 MKMLKLTDTKSDLLLQNSYYFSQCKLLKYSLMLTGAWPFMKKRIRNFLHAYQLIMMS VLFLGQLKYLYQHRNNTNELCLNALWLSLTSCVIGLVLHNMLFNDYVFAMLRAVVSV YEIAKSDKKYFLTMIPLSMIRKIIAGFYVFLCISSVVGPILLGMLRCDMKSPDRFQCMKKY

438 SIFKTTFSYKIYLISFTSSVIFCTLNIYHFYDSVANMLVVVITAKLTFANIISNIKYETHDKSI TKNSDYVNLKSTTLRILKRHQDVIGMIDLANDGYWMPILICISINIFTLAINLYTILTFIDKP KIVLTVLPIFILCLGFPFIVCTGGQQVLELNNSLKYAFYSSQWMDFPKDIQTTFHIFQVRC NNNLKICSCGLIVVTTEFYVNMMRASCSYFTFLWSVKHKS >TbOr95 MITKSNKNSIIIKDTSLENSYYFNQCKFLKYSLILTGAWPYMKIQIRNCLHAYQIIMIFVLY SGQWKYLYQHRHDIDELCLNALWISVTSSMIVLILHNKFFGNHVANLLRIIVHEYEINNS NKQSLLYMIPLSMVRKIIAGTYFAVFILSFMGPIILGLTRCDMKSLDRFHCMKKYCISKTT FSYRIHLISFTSSAFFVFMNVFHFYDSIANSLIVVISLKLNLENDLMNITNETDYGSEVINY DYVSFMSEARKCIKKHKFIIDIIDMANECYWMPILLCISIHILMVSLNFYIISISFDKPQKYV SVLPILFVCFVFALIACTAGEAVSDINDSLKYAFYTSLWMDFPKNVQTIFQILQLRCNNS MYITAFGLIVLSMEFYVNTLRMSGSYLTFLWSVRD >TbOr96 MKKNETEITKGIFDTHYHLVKRMSIMIGFWPTQSRKVRCFLHTYQMTTSFLILYGLSSYV LQNKNQYDKMFTNAIFFFIHVAIIMFLANFISKGRSVNSLIDMMDKDYKIYMRNKETQA FFTKKFILARKMMLFSLAVIFSFIIFAVVAPIVIAIFRNTFKESAIMYDTVFTFWFDTACEIS SMTYSHFTIQHFCSVFVIGVGSIFFKFYSMRFHLQDYGKNYNENTGYSGNPYIIELIRTVI KEHWELFYLLRLVNDVIKLILLCGVFVMSSLIVMTSLVIKISMDNPMKALPLVPVITFSV GGLGLICFIGNGVSNIPNEIRLAIYKSAWIRVSPKINKSLIIISMKCFYPCELNAFGLIIPSME LYISLLRACFTYITVLLTLQEKDN >TbOr97 MKIYGFHLTHINVKNYDNVEFNQKCNTDIFNTDYYHYIKSVSYYFGLWFYPSTKMNCL RWAYFCMYRCLELMVFSCQVKYALHFLDDLDELVSCLVWSGILFLFFSAIIGMYVRHH CTIALEDIQSLWAATTDPEERKILHDNAEMPRFIIACIAVAGNTMLFMNIIASIPPLMYEIIT SKHTVYNDFYLMKFEILFETKYDMYAKKFIYTCAGFQFVFGFQLFLGLCFIIMYHLFARF SILRSIITKDITQTTDNLVNGKKTLKRFIMFHADSLKYFYHLNEMAKDNLLMGIVLNVST VTIAVYTINVNIGHIEKILPVLPMFVSLMIVIFLISYISEEIVQMSTALVQNCYHSPWLKWD VNTRRTLNIIQCRCSIPCEVSAYGIVKMSLQYYTKVIQISYTYLTFMASMRTK >TbOr98 MVSGKIRSTHGFAMHEFIRIGHLLGIWPLRNVLEDKMKQILFISFNMSLVFGQIRRVMISK LHFREMCFSLLWTYPNLLSLVLYINTIARRKELSRTIDIVTNDYNCQLENYTERIMKNDK FTKLCIVCLYITGCSGVIFILANIFLFKSDSFSLVVAPKFLAKYNRELESGLWATMLFLFIH QIYFYSCSGLTAWNYPSAKVDTIMSFCFRKRFLSIGPDPDKIKSLLVIKKCSLETCELLFY VKLLNDICAFLLLLGIPAAIFLSSFSLFLLSISVDSPDIWVLFSSITFIVYPNVVLGFISFFGEN IQSGSGRLKLAIYSINWCDPTKIDGKSINIMQYICNRPMIMSAYGIVTTTLENFVQVLKVT FTYYTLLLSLES >TbOr99 MSSQHKGIIGPRKFPNLDNFAMREFERVGLLMGIWPLRKLFEDRAKQIFVILFYLILVSGQ IQRLMIAQLNFRETCHSILWIYPNVLTLVLYVNTMSRMKQLTRTIDIVNSDYLYYLEDYT KVIIKNDKFAKFCFVSLFTSAIIAIIFITFHVFVLRTEHFSIVVPPKSVANYNKQITMSLWMI SVFYLVHQIYYYCCSALSGWNYPGAKANAIIKQLMSISMECDKIDSLLIIKKCTAESCKLI KYVHLLNDVCAFILLLGIPTGVFLTSFSLFNFSPRSPDISIIFAIYPNLMTGFMCYLGDGIHS GSTRLKFAVYSLNWSDGTKVDIKALQIMQINFDQPMVISAYGIITMTLENFVQILKISITY YTVLLSLDAEK >TbOr100

439 MSIQTAGDLSMVEAEKVGQLLGIWPFKNHFDDRVKQVIILMYCLIFFRGQINLILNLQLD FRDLCFNIMWIYPNFLMAFSYLNIIHRRKQFVEAMNIITHDYVNHFEDYKKIIMLNSKLT TICKIGIILAATSMYIISLIPLVFIRFIKTDMLFTPLSETSYNKITHTILWSIYIVLFSNQIYLYC CLTICGWNYPVAKVDTILKKIFSINKSVDKNSTSKIIKSCGADSRVLSQFIKLLNISASNML FIGISASVLLTSFSLFNLSSKLNDLSKLFSSIVFIVHPNVMIAALCYFGDNVSCLSAKLSYG VYCINWPDKEKSEAKLLNIMQILYNKPIVVSVYGLMPASLALFTQTLKISFSYYTVLKSM EI >TbOr101 MLGIWPFRNILEDKLKQIIFFTYSSTFIRGQINFMLISKLGFRDTCFYVLCVYPTFLMAFAY INTIHRRKQLIEAIHIITQDYVNHFEDYKKIIMQNTKFIKICIVGIILASISMYTLSLIPLFFLK YIKINIEFSPLSATSYNKNTHAILWTMYVVLFANQIYFYCCLTICGWSYPVAKADSIVKKI LSIDRSDDTTSTLSIIKICSAESHVLSNFLKLMNIIASNMLLISVSASVFLTSFSLFNLSLKLT DLSKFFTSIVFIVNSNVMLGFLCYFGENVSSGSYKLTHAVYSVNWSEKGRDDAKALNIM QVLCNRPIVVSVYGLMPTSLALFGQSLKISFSYYTVLKSMES >TbOr102 MRLLIALKEFFVLLQAKTVWPEDNNVFITYFRHYVLLSKVIGVWPYQKLVQHIILIGSYA SFSDAFFISQVLYLIKHKNDFDNYIGNFIFIFAHVFIYVTRLNLIFYYKSYLKCLMEYNAD YEKSKLNESEGIIFLKTILFTQKILRRINYVFAFTLILCTFWPLWPKFMDVINPQAEPRQEY YMIDLNIIFSNLDSFTMKYLYTEIAIMIGFVHFQIIIEYSVITFYHLYSRFLILEKKLNDFRQ YEDRVLPKDTFNKKIGLCVIRHESCFELSLYFNNLIRNIVLFGFTVDMMAISITMFYIQMN LDNIIRILPKLLPLLGFPLGLVFALCYMGENLRQHSIHICETIYNLPWFVWKSSINRSLDIM KFRTLRPCEVSAYRLVTLNFEFFLRVFQTSMSCFTFLLNMKE >TbOr103 MIMKVTTVCGKLYAVLKYRAFIAQDEEIISMYFRLYVLLAEIVGIWPEQTVSQQLLRTAC LGLLSIILLISQVLYLAEHKNDFDNFTGNFIFIIAHMFIYVTRLNLIIHYKKFSICLAEYNKD YKKRKTCESEGNIFYEHILKTQRILQLVTYAFIITLILCTIMPLWPKILDAINPLTEPRPENM MIDVNVYFSTSVSAAFIRKYLYAEVGIILGFIHFQIIIGFCLLIFQHIYSRFFILEYKLNDLVK YENRILAQETFNNKIRLCVKQHKSCFELTTHFNKLIRNIILFGFTIDMMAISITMFYIQMNL DNLVRILPKMIPLLGFPLVLVFALCYLGENLRLHSIQICDTIYNLPWFLWKSSINRNLDIM KFRSLRPCEISAYGLLTLNIEFFVKIFQTSMSYFTFLLNFKE >TbOr104 MLDPKWDDNETESFFNTERDKVYNSYLKIYRIVLVALGYWRRQSKLVSFLSRLGQSFIVI LLINGQVLYVIENQDNYDEVCTEMIWIGCQCFVFVNLINNSLYKPMIFRGMGLYQRSFD VYCNDDDSLKILRQDSLVLQKYLKFSAVILQSFWLILYVSPFVLAISTNTYDMHTIMNDY QLVVKTAYPPWVDFILLYASVFSTGLSVQLFTCTCVSGFGAIFLQSKIVQNNLKKIGELSE SSAKSLVAFCVKEHLIVFEFVYLINEMAKLLIVFGFMIDLVELSLSVYNIKTNSDDIGKILV TIPIPLFVYGAPGVLCMVGENIRQTQDEISMAIYQSSWMKLPPKIIKSLIIMRIRCTVPCEV SAFGILTLSMELYIKVVRTSFTYFTMLWTMRENNMS >TbOr105 MIHQESVILQKYLELSAIIIQSIWFIFNLSPFILAICMRIYNMNRIVHECLIIVKTAYPPWVDF CLLYFSSTCTILSLQLFTCTCIAGFGGVFLSSKVLQTKLKKIQSGDSKKSSAKSVVAFCVR EHWTVFRLVGTINETAKLLIIFGFLIDLVELVLSVYNIKMNSNDFKALFVTLPIPFCVYVIP GVLCSLGEDIREMQYEISTAIYDSSWMTFPQTITKSLIIMRSRCTIPCQVSAFGIITLSMELY IKVIRTSLTYFTM >TbOr106

440 MSEKKCENDCTVQVTVAKSKRGYNIYLKVYKACLLILGYWKTQNKFWSYFSQTVQLV LCIVVVYGQILYIIENRKDYDVVCMEIIWMCIQIFVLVNIINMNINKHLIFRALELFQRNLD LFKDDYDTMKIIQQNSLLVKKYLELIARIFQSCWLLLNLFPFVLALCTKVYDIKHIMREY QLVVKTTYPDWVDFLLSYPSTTFTILSLQLFTCTCISGLGAIFMQSNVVQHKLRKLGSED CGRNSTKSMVAFCVKEHLIVFEYVFIINEVAKLLIVFGFLIDLVELSLSVYNLKTNTDDIK TILVTIPIPLFVYGVPGILCLVGEDIRETQNEISMAIYQSSWMKLPPTITNSLRIIRLRCTVPC KVSAFGIIILSMELYIKVLRTSFTYFTMLWTMRENNMS >TbOr107 MLERNCENDPKEIAAKRDKEFDSYIKVYRAILLVLGYWENQNKFLSYFTQLVQLTICLV VINGQINFVLDHQDDYDEVCMEVIWICIQMFVLVNIINTTINKHLIYRGMVILRRHLDTYS DDKDSLKIYRQNLSLFQKYLQLSAAILQTFWLLLYLSPFILAISTKVWDMNRIMSEYQLI VKTSYPPWVDFILLYLSTTSTILFLQLFACSCVTGFGLVFLQSNIVQNELQKLGSGTKTPA KSVVAFCVKEHLVLFEFVYIINEIAKLLIVFGFLIDLVELSLSVYSIKTNADDIKTILVTIPIP LFVYGAPGLLCFVGEDIRQTQDKISMAIYQSAWMKLPRNITKSLSIIRLRCTVPCEVSAFG LITLSTELYIKVIKTSFTYFTMLWTMRERNMS >TbOr108 MFDSYFRHYKIITKITGCWSYQNKNFCLMLHAYNVSVAVIVFYSQVAYLYAILDDFRNV CTTCNWSFYTLLVIFIMFHNMIKKKKIDRIINVIDNHCKFILQNKKHEKDILKQLMITENV SKIITRIFTVLWILVLVLSSAAMLYICHGSVDKQLCIEKLLILQTTLSNKMAILITFTNAHFI AYYVNGFVLFCNLVLCILNIEISVVQNELMSLNTRKQHLYNNTNRTSLRFSVINHQFLLK FMVFVIEIVDLIFLFLCIISITLITSSIYTIYYMMASSDYNTLIIIVPCTIADLIITLTFCHYGET VIEANDKIKIAIYNSNWTMLGPGMKSELKIFQRRCNIPVVINFYGVITLSIELYVKILKTCM SYYTFLGAVNYE >TbOr109 MLHNIIKKNKIDRIIAMSEKYCRFFLQNEKYRDEVSKKLQYSESLSKIITSGFTIMWISLPF LSITGMLLFCHSATDKQYCIKKFVILKTTLSNKWAIIFMILNAHYYSYYGNGFLLFCIIILC TGYTKISVIENELNSIDENMQQFDSNVNRKSLRFSIINHQSLHKFGTSLLEIVDLITFLLFTI SISLLTFSVYTIYFAMESNDFDTLLIIVPIMFQDLILTIIFCELGQQSIDINDKLKISVYGSKW TVLSPDVKQNLKIFHRRCNIPIEINFRGIITLSLELFIKILKSSMSYYAFLKTVNHH >TbOr110 MRLSPAMRENKKSSTLVEENLFDSYFRYYKIVSMITGSWAHQNKRACVLLHVYHGLSA FMIFYLQISYLYSIRNDIKNVCTTCQWTFITICIIFIMFHNMINKKKINIMIKTVENYCKFIL QLQKHKKVALQQLKINEKLSKLITRLFTIMWISLPFLSLAGIVFFCQNTTDKNLCIKKFIIL KSVLSNEWAIILTISSAHFFSYYGNGLLLFCTLMLCTRNINISVISKELESIDEDVQYLNNR GHRTNLRFSLTNYRFLLKFFSFLIEIVDLITILLLTISITLLTFSIYTIYFSVESTDYDTLLVIVP IMLVDLVLTVTFCRLGQMTADTNDQIKTLIYGSTWTVLSPDMKLNLKIFHWRCNIPLIIN FRGIITLSMELFIKILKTSLSYYAFLKTINHH >TbOr111 RILTTINDYFEDAIRNHKDIVLKHLIVCENVTKLIAKFFTLCIIIYVALIFISIYIFCYARKCA DKESCVKKFIYLKGTFSNEWMISLIVLGVSPVFYTVAGFVIFSILALFTGYTAQSVVEYKL NEISNSDNFVCTANERKFANTIRYCITIHQYGLELAKLINDVIHIIVLVGLSLAAGGVALSI YTINSLWKDSNHDKILSIAPLFIMELILLLFFSVIGEEVTCCGKIVKFFIYNTSWTTPNAKA KQNLIIFQMKTDIPVVIYVFGNLCLSLNTFVIIIKTSMSYYTFLKRIS >TbOr112 MIEEKEVKSIDKSIFDTYLRYYKYVSLFTGFWTYQNKHICLFIHSYQLVLLTAYLYTQTV YLYTIRNEFEKLCIDSAWYCVILFIGTNFVNNIVHKKKIDRIINTVDDYYKHLLRVYKEIV

441 LKQLTMCENVTKTITNFFTVAMLIVHFLCLFLIFTLCFSKSGNEKDMCIKKFTVMKGTFP NEWMIFLTFFNATFSTYTTASLVIFSVIALCTAYTKVSIIQKKLGDVSDLGDCLSTADEHR FVKNIRFCIIEHQSALEFAAFINEVLHVIAFAGLLLSVGGVAVAIYTINILWKDSNHDTLTS ITILFGLELLLLLLFNAIGDEVTYLSGALAKAMIYNTSWTVCSKPKQNLIMFQMRCDVPI VIYVFGDIYLSMNTFVNIIKTSVSYSTFLKEMG >TbOr113 MEVIKTVETTYSYVVIKKYQPIYICYEYFFLICFTGQQLTEKSSQFGKLFLTTKWYNLTKE KLFIFSLINVYGQIPAYITAGKMIKLTMETYTDFGRVVNMIDEHYKYAMRMYKEIALKQ LTLCENSTKTITKFFTIVMIGILLLIFILMYTFCFSKKSTERDLCIKKFMLAKGTFSNEWMIL LTVLGASFACCSAAGLVIFSIIALFTAYTNKSMGEYKLNAISNFGDSVSAEDEHKFTQNIK YCIIIHQFALKFASLINEVLHLIAFVGLLVSVGGVAVSIYTINILWKDSNHDTLMTIIILFCM ELILLLLFNAIGDEVAYLSGTLIKFLIYNTSWTTSCSKSKHNFVIFQMRCDTPIVIYVFGDII LSMNTFVKIIKTSVSYYTFLKQIT >TbOr114 MTKEKGVNSIDTSVFNSHFRYYKLLTTPTGFWTYQNELIRKFLHVYQTVCFCSLAYTEIV YVYGLRHDFIQFCVDSSWLFIGMFVWAIFFVFTVYKKKVDRIINFIDHMYKNALKSHEQI AYNHLTFSRLLLMVIMYLCKLFLLILLPPSFLLMYIFCHMRKSDDKNLCIKRFLVIKGSFS NKVMIPLNVLCLYFYIHIVSGFVLFELILLTTAFTKMTIIKYKLSEISHIEKFITIAEEKEISQ KIREIINSSLSGIRFGTAINEVIQIVAIIGTNACVCIISLDIYSISLQMENSNYNTLIFIIPILWV ALAHMFGSCRMGDFAISMSGKDVQNAIYGTAWFMFPSNIKKTLLILQMICDNPIKLRIFG NLILSTETYVNILKTSISYYTFLTTIK >TbOr115 MKKVQKVHSISESIFYTHFKYYRIVSTSLGFWTYQNKFFRKLLRFYEAVLFSAMSYTQTR YLYGISDDFVKFCVDSSFYWVLLFVWAIYFINVTQDEKIDRLINFIEYTYKNALRTHKEIA CKHLTLCEKFAKKLTMTFRLMRSLMSALIILSICLFCHNTEKADKNLCVKKFLVIKGTFP NNMMIVLNVLSIFASVYIVASFVLLTISIMTTAYTKISIIEHTLDDISRIDKFVNNTDKNEIE KCVIKSFKDYQSCNELASIVNEIVQIVAIVGLLVNIGLISLTIYTIHYVNSNHNALFFILPIFL VTMFDLLIFCRMGDEVIILSGKNMQISIYNTSWMMFPSNTKKILLIFQMRCDKPMVLRVY GSLIISMDTFIRIMKASVSYYTFLKEMA >TbOr116 MTREKKVNLIDKGVFHTYLKHYKLVTSAIGFWSRQNKLICRLLNSCAGVCIVVMGTLQT IYLYGIRHDFAKFCIDSIRMIEASFVWAIYLTYMTQGRKIDRIINCIENRYKIELQTHKGIA LQQLTKCEHIIQILAKILKLITCSLISSLFLSVYVYCYTKEFAVRNFCIKKFSFVKSTLPNEW VFVLNCLSIYFSVYIVGSFALFSIISIYLAYTHISIIEYRLDKISDIEESVALGDEHKFGKNIK MAINVSQFSLRFGDMVNEVVQIITVVGLFASIGIISITILAINYSLKNSNYDSTFLIIPVFFV MDLTLLLLFCQMGEQVINISGQHQKLRLYQTSWTMASCKIRRMILLFQMKSEEPIVIRIF GSLSLSMEMFVNIIKTSVSYSTFLKEMG >TbOr117 MTKAKDVHLINKSIFCTHLKYYKIVTSYTGFWTYQNKFIRKLLRLYHTICIYLISHAQIVY LYGIRQDFVKFCIDSSMFCGALFVWSILLTNLVQGKKIDRIVNLVEDTYEYAFQMNYEIT CKQIKMCENITKSVANSFTLIIYLLLSSLITFIYIFCRGRESNDKTLCIRKCLLMKGTFPNG WMIVLNILSIYSYMYIVSGFALFAIISVFTAHIKKAIAVHKMDKISNLDKFVQKTDEHKIE ENIRITLNLYKSALKFGFSINEIVQMISFVGLAVSIGIISLTIYTINYLLKMSNYGTLMFIIPIF LVNLCLLLLFCGMGDEIIIMSSKDLKKSIYDTSWTMLPSNARKALMIFQIRCDTPIVLRVF GSLILSMETFINIIKTSVSYYTFLKEMG >TbOr118

442 MIKEKKVHRIDRSVFYTHLNYYRLATSYTGFWTHQNKYISILLRSCQAVSFCFLSYTQSL YMYEIRHDFVKLCIESSLYCVTVYVWSIFLSNIIQERKINRIVNLIENIYEHALNVHKELAY KQLILCEEIMKYVTNCFMLIGCLILLSLALFIYIFCYKRKCDDKDLCIKHSLLLEGTFSTK WMIVLNVINTYFFAWVVCGFALFSIISIFAVFTNKSIMESKMDDISHLEEFASTAQDQKIT KTIRESINFCKSAIEYGYLINEIVQIVAFVGLAVSIGIISLAIYTINYLLKSSNYSALNLITPLF FCDFCLLLLFCEMGDEVVMMSSRELKQTIYKTSWTMFSISNRKTLIIFQMRCDVPMVLR VFGSMILSMETFGSILKTSVSYYTFLKEMS >TbOr119 MKKGNRVQLSNKSVFHTYLKYYRIVTSSTGFWSYQNIFIRKLMHSYQAICLCLMSYTQT SYLYGIRHDFVQFCKDSTWFCAAFFVWAIFWINTTKEKKIIRVLNMIEETYEHALLEHSEI AYEKLTMCEDITRKVTNLFKLIIFSLTVSLFLFIYIFCYKRKSTDKNLCIKKTLLFKGTFSN EWMIVLNVLNTYFYMYIIAGFALFGIISIFTAYTKKSIMEYKMGEISDMDKFISNTDEQKV ASTIRKTIIIYQSSTEFSVLVNEIVQIIAFIGLAVSIGIISITIYTINYLLKNSNYDTLFLIIPLFL VVLCLLLLFSEMGDEVITLSGIEIKLCIYKSSWTMFPSNTRKNLIIFQMRCDTPIVLRVFGS LILSMETFVNIIKTSVSYYTFLKEMG >TbOr120 MIRKKNIHLIDQSIFNTHLRNYMIVTSCTGFWTYQNKFIRKLLHSFQAVCLTFMSYTQTA YLYRIRHDFVKFCVDSTWFCVALFAWAIFLTNATQSKKVDRVVNLIEDTYKNALQIHSEI AHKQLKASEDVMKVVTVLFTLMVFVLLSSLFLFICIFCYYRECDDKNLCIKNFLLVKGIF PNKWMIILNVFITYIYLYIVAGFALFGIMSLFTAYTKKSIMEYEMNKISDIKEYVDITKEH KIAKDIRRTIILYQSATEFSFLINEIVQIIAFVGLVASIGIISITIYTINYLLKSSIYDPLVFIIPLF LADLGLLMLFCEMGDEVIVLSGHEIKLSIYRISWTMLSSNIRRTLVVFQMRCDTPIILRVF GSLILSMETFVKIIKTSVSYYTCLKEMS >TbOr121 MTREKEVHFVRTSVFYTHLKYYRIVTSSTGFWTHQNKFIHKLLHSCQAVYLCLMLYTQI DRYVYLIEDTYKHALQVQKELAYKQLIMCEDITKAVANLFVLMICLLLSLLFLLIYIFCYI RKCVNKSLCIKQFLLVKGSFSNKWMIVLNVVTIYFYMLLVAGFALFSIMAIVTAYTKKSI MEYKLGEISDLEELKCVSSADKHRVTKTIRDSIHVYESATELAFLINEVFPIIEFMGFAVSI SIISLTIYNVNYLLKNSNYDSLIIIIIIFLVDLSLLLLFCEIGEEVIILSGTEMKFVIYNSSWIMF SPYNKKIIILFQMRCDTPLVLRVFGNIILSMEMFVNMNYVDETKTHKILTDEININNVTIL >TbOr122 MHGRQFNSIYDTHIYKEYKFVLQLNGIWPDQPFYAKFFIVLITRGAFNIFMVYKFHYTISL FGHSSKFVPAFVDTFSLCLFYTVDLSMLLQRKKYLKMHYDIENDWKNSLYDSNDKKTF NEHSLKCKAIMSKITKVLGYTCLITNIAPCIPIVLDLVSPLEKKRHRVYLFTTVLTEQLAN YYFVVYIFFDIVFTILMLEICVLFIILAVLGFHAAVLFAILKSNLTNFGKSETKNEIVSEND MSYIHNQLKRCLKQHWNALRFFNLGESYYKPLLLIGFFQNVSGTVACFYALRKNLKNW QIMLRGIDLVLFNLVYCCIGEYVLNFTEDLKIACYNSNWINLPTRMKKLLSIIHCRCNKV CVVSAYGVFYAKREYHLKFLQISLSYFTVICSLTDRKSV >TbOr123 MRKTLTSRYRKEGVIFDNEFHKIFKILNITNGYWFRYPTWLNIIIIACNRVPLVLLFYYEM VYIMEGNEDIELVPKLFDISILTILYVVDFNNLFKRNVLIYLLFDIINDNSKVLTGFRELKV YRDFCAKSQKVMVNVTRLFIITFFISTSTPLISKLVMYCIYKENRPVSHITTSSLNTRFAED FFLLHFIYSICAMVAINQIMIIFIFQVVLLHHFSIVMIISGFTLQHLGEIDPFHNRNDLDQNK AEHDMRCSLKYQMDCYKIFEATQNFVQVPLMFCFFQNAVALSACLYQLTVTKDDLNET IANIIRIIDLVFSNFCYCALGQYCLNKFELLLEESYECSWFNLSEQNKQIFYIMRCRYMKS CNISACGLFNASLESHVKFLQMSFTYFTLMTTFQ

443 >TbOr124 MRQTLTSRYRKDSVIFENDFNKIFRMLKITNGYWFKNPVWLNIFIIILNRIPVTVGIYYEM VYICEADSKDDAISGVFEISIFSLLYVIDLNNLCKRDVLIFIMFDVINDDSKVFTGCREKQI YDEIWKRGQIMISNIARLFFLSLSAYLMLPSISKITLYCIYKKNLPVSHLMTTSLNTRFEEN FLLLHLIYSVIYITVLNQVLVIMIFQIVMLCHFTTLMTITGLTLKHLGETNDIKNMESIDQN KAKNDIRRSVKQQLKCYRMFEYVQNLVKLPLMVCFFQEAVALSVCLYQMVFYKNNFH ETISRFLKIMSLVFTNFCYCFIGQFCLNQFELLLEHSYESLWYEMSKHNSQVFHIMRCRY MKPCNISACGLFNASLESHVKVFFSIVSLIYVNIVVIIFFQFLQVSFTYYTLLTTLN >TbOr125 MGESFFDTSTFEVYRFMTHLVYQWPFQKWSQIFLVMSVEIPISMMIIYGQLTSVFSLRNQ KDEMIISSLWAVPHFYIYIININALMNRKITRKYFQEMKADYLTDCIEDYNVLKPYGDNT TKLIRLILMSYVFCLIPWISGPLLAIYLSEEPVDIHSDLYFVFDIDLERLSKSLFLKCFAYFY GILATAFNYSTFSNTCIIYFFHLFGKCAILKSRIENMGKDGNSHPGTSIIRDIKNCAKIHGK LLEYLEILQKWGQFLLFVGVIINIFVISLALYTIQVNTSTFTLMRYSSVYMFIIFSMLVLCIF GERVLDINVQIKQSCYDSDWLSLPKPAAKIFNMMQCRCNRNFGIHAYNLILLSMHYYVK VAQTAFTYFTLLCTMTNK >TbOr126 MKMYSDKDYDLFYKKSQYFYCQRILNILGGHWVYQRNWERHIVHGLQLLAVFSLGVM HVVYIKIHLGNKQNVFTAMLMMSCCVIVLFNSALMHLNFKKIKHIYDEVALDWITVKC QIDKKLMMNEANAFKKSVTSIGAGLFFILFLLMIAPMARFLINRYITKNNIVPVYYFELYC CHKYAVVLKYIQYALGILLQNGNFYVSISNIFLMFSYIYSKFDIIESRLERIDDRNNEATTY VNLCVIVKLHQKIIKFFYLMLDTLSMLLFVGVIYNVFLMIFIFYFIKLNLNDLGRLYKVAP LCFCAIIVMYIFNAYGQSSIQHSLQINSACYNLCWMSMSSNETKIVHIMLCRSQTPCVLSS YGLITLSKESYMKALQTAFSYFTVLWTVTGN >TbOr127 MYKLETMKSSDEEYDLFYQESHYYYIHRRWCLIAGHWIYQKSWKRFILKGSQFVIVLIM TLSHMLYMKVHWGEYKNVFTATLMMSSCTIIIIIFIFCEQNFKQIRCLYDEMALDWITIKC HNDIHLMLNGADKFIKLATKIGAGFIWINVMMMVAPIARAIILRYIITNNERVPIYYFEILC CLKYEAIIRYIQYTSSVIVQFLHFYVLISNVFLVFTYIYSKFDIIQCRLESIDCRNSIATTYM DICVIVKLHQKIYKFFSFVNESMKWLLFFGVFYNVLLMCFIFYFIKLNYNDAARMYKVIP IGFIIIVIVFVLCIYGENMTQHSLQINNACYNMCWMNIRINEQKTIFFMLCRSIIPCKISFFG LVTLSNEYYMKVMQAAFSYFTVLCTVAD >TbOr128 MFITFFCHLRHKMFCPWWNSTRKQNNCKVKCNKDKFRCKGAVNERESIFDKYFCIQKY LCYAVFDWPAQRVLYKNTGNCFYLIVSVYVLSCQINYILQLQTIDEQISSFTWVICHLFMI TLFTNNLYTRKIMKQFYELMDDNFSYFKAKKDREIMLRIIKLGRRFTQITLFVQLSGGVT LSMLPLLVGIFSSSQRENYFFVQLKLIFINNFYLKCFLHGFVIIYCVLYILQLDNMIIISCVN MFAFLSILEYRLKRLGNFCKDSEYFEEYTINELKFCAYQHQNFITSISLINEWCKFLLFVG LSIGVAEISLCIFFLVKNLNDISSIIQLGPILIFSFIGIFFICFACDHIQQTNIKLKHYCYESSWI NLSTKSAKMLCMILARCNVNCEITLSGLFTMTLEFYVKMLQASFTYFTLIMSFNEQT >TbOr129 MRYEYNNVIVKQRNWTDKEVQSHLKSTETGVGSIFSTRHYRIYKKFDIITIKWPFLNPYK IFIARFMEFSFSAYFLCCQVSYIKNLPDLDSVIVSSLYVAIHIFVLSLMVNNAMKTKSTRL LYEMLDNDVHFYDEKEESKIIRRFHVIVGKVTGYLITFHVSAGVVVSIAPSIVYLQPSIGIN LFASDFEIVLLKNSYLAKCIVNMLSIMETLNYVLVLDNIVVLIAAFGFAKFTIVQNRLNRL GKLEEGQTKVNLATANDDIRFCAYVHQNLIEVHDLMNDYVKIVLQSGAFLSVGIMSLSC

444 YCVLKNLDNPTMILKISPVLIFLFVGLFIFCKLGETMTESYENIQNWCYNCEWLNLSPICI RQLIMIISRTNTKCVIKYYGLISLNMDYYVRVLQTAFTYFTVVTSLTE >TbOr130 MHWEYPNMFMKRQEWSENQTKLNIDKLRTGVNSIYETRYKYLKLACIVSFLWPYQKLS YKIVLRTLQFFLACYIFFGQIDYVLSLKTLNDIVTNVVFSGVIIFAMIVFTHFWVYGKEQK QIYEIIENDWDYFEDEEDLTILNHYTRYSAAIIKWLTIFIPIAAFIGTMAPVYGLYIPMFYN LFAFQFNLLWLEQDNYNFKCFIYVIGVTWIVMYIVVFDNIVALISFQGYTKLIIIKNRIKK VGVVKDKTGSYDAEAEKHIRFCSLLHQNYLDGFFVSIEWGQILMVFGVLIGILELSLAFF YISENMSDFVNLVKIIPVVFFAVSCLFLICLSGDIMTENSFKLADICYLCGWTNLSQRNLK MLSIFFMRCNFKCQFDFSGIVVLSLQLYINIIHASFTYFTVIITVRRSEERRVGKECRSRWS PY >TbOr131 MSWEYSKMFLKRKEWIENRPKMDVNKIKTGNGPIYEGPFKHVKMISKCLFLWPYQKM VYKIIAWIMELSLTLYMFVGQLDYISSLKTLDDIVTNGVYSASFVFVVVLIIHYFTFGKEQ RMIFEFLENDMYLYQDEEDFAILNSNVRYSSLITKWLTTVFPMATIIILILSIPFASYFGVL HKVFIFQFNIVWIKNKSYAFKYTTYSIFVVVLIIFINVFANFLIIISCEGFTKLSVIKNRIQKI GVAKDGQKDICQLESGKQLRYCCYLYQDYYVTFITITKWGNILMVVGVFIAIIGLSLSLF SVTENLNNFLNILKVIPIIIFTLPCLFFLCLAGDHLTDSIFEMADSCYFCGWTSLSRRNLKIL SIFYMRCNIRLQITFAGIVTLSMEMFVKILQSSFTYFTIIITMRDKTLN >TbOr132 MDWENTNLVLKKKESSESKTTTVDKVKTGNGPMFEGPYKYVKLTSKILFLWPYQKTIY KIIARTFQLSLTTYMFICQVNYLLSLKTMNDVVMNAVYSISFVFILVLFVHFCVYEKEQK MIYEYVEADLDIYQDEEDIATMNSYMKYSCLITKWGYIVIPTATALVIIITPVYGLFVPFY KVFIFQFNMVWIRGNNYTLKYIFYGTFGWTLVIHVLVFGNILVVLSYEGFIKLSIIRTRIQK IGSVKDEQTDLCNVESGKQLRFCIFLYQTYLESFGIVVRWGQILMVFGVLVGIIGLSMAF FYISENINNPVNLLKIVPIIAFAVPCLFFLCFVGDHVTESSFKIAESCYLCGWTNLSQHNIKI FSIFFMRCNIKLQFTFSGIVTLSLQMFVKIVQASFTYFTIIITLRDKNLN >TbOr133 MSWDNANLFWKRKEWSKDQQKLNIEKIKNGNGPIYADPYKYVKWNCQVLFLWPYQK TTYAVIARTIHILFTVYIFIGQIDYILSVKTLDEFIRNSIFTASFIFLLVLFVHFCVYRKEQKK LYEYVEYDMDIFQDEEDFAIMSSFLKYSSLVIKFLSIIIVSGTILVLIITPMYGLFFPMTYNV FIFQFNIVGFGKNSFALKYTLYAIFCFTLIVHMLIFGNLLIVLSVEGFTKLSVVRSRIKKIGL LKDGQPDLCDISCKQNLRFCILLFENYLASFVVVTRWGQILMVVGVLIGILGQSLTLFYV TQNFEDILGVLKVMPIIIFAVVCLFSLCWAGDRLTENSLKIADSCYLCGWTHLSRNNFKIL SIFFMRCNIKIQFTFYGIVTLSLQMYVKIVQASFTYFTIMITLREKNFNLQPTG >TbOr134 MSSSPELYYKTTFLEDQSIFESKYYKSYKNISLWAGFWPYQNFKARIFIVIIHLLLSFPTIV LQFQYIQAYAHALDEKINSLFMIGVHSFVYIFRFNKILYLNEYFLLHDEINKDWMYAKSN ELRILHIYTSKARSQLRYAYNSTFSFYIIIVLTSIIPTMQNILFPQSERRKEQYLFNTALNIW FADYYFYLYGYYCIIVTMGTLYILLFYNLLYMMTFHTGACFAIIALKFQSIESLFNIETIYN QIPKNECNKCFKQHIKAIKFADLFCHVLEPLLFISVFITLMCTASVIYATQLYTSTGNTNE MFRWIPLVICTLNTSFIYCYLGECLIESSSNLRNERSLNIMQCRSMKNSELTAYGVFVSSL EAFVMILQSSFSYVTVMSTFDK >TbOr135 MHCQTESVFDTHYKIYKYSRYMNGVGFVQPLYSRIFFHFFNRLFLTILFILEGRYLYFNR RELQKISQNIYDAGNLVVVYFFDVTAYVNRHTIHRMHENMKIHWNSIDSYDEYIIFYNY

445 WMKGKKYLTAITGLFSFLYIFLVLISPVTLKIFDSLKPLNESRRECYIVETEINSIYSDKFW EVHLFYVFASTAAILRMHLIFCSILVLTLHSLSLTDTTGYYANQIGKNVHNELSFKKNAIE DIKICYKKHHNAEEFMNGIKQVFESILFICFSISICTLSTTLYTVRKLVQEKIDLSNALELIK VVLLTTSIYVTNFMYCFVGEQISSHFEELNSKCYDSAWYKSSPRIQKLVGLILCNCKKVR CVEAYGIYKASMDCHVMLLNTGFSYFMVLSSL >TbOr136 MHCQTESVFDTHYKIYKYSRYMNGVGFVQPLYSRIFFHFFNRLFLTILFILEGRYLYFNR RELQKISQNIYDAGNLVVVYFFDVTAYVNRHTIDRICKDMENQWAEINSEEEYNIFDKY SNHGKKALVCLTAMFSIALFIGNLSPFVPLVLDFFLPLDEPRPQTHVIETEINYRYKNKFL QVYVFYFISVMFFIGKMHLIFSFLIVTPVHVMTLAESASHYIQQLGSDYYILYGKDYQYIQ KSNANEIKFCVKKQREADNYIDCMKQVYELVLLLCFIISICTLATGLYTIVITISEKSDSSN LIVVLSTIFFIYCANLMYCTVGEKIINHFANFNYACYCSKWYLLPLQLQKSLLMISRHSQ KLRSFNAYGICTASLEFHLQFLSKGFSYFMMLCSLSD >TbOr137 MNIFESDYLKLYKIDCMIAGTWPHQTIYTAVFILSLQGFISGFLSITQIRFIHNFWDKKEIV FMAILFTLIHYMMVITRFHQIIFYQRIKRIYAIISKDWKVITPGTFEYDIMINYAVKIQRMIR NCAVVITTNLMAFYILPLISKLCDILYPLDTPRPDFLLLNMQLKLIKRHTYAAKYIMNVSG LTTAFVLIMVSISLMTLFTEHLFMRFVIARTRIEHLKKFDTHSDEKMAHFSIRGCVSYHDS TLKFAELINKCLQLLIVIVIAITTVILTLTLYNIFTVPNITNFICKMLPVSFYVTMTIFYFCHT SEKVITCSTDVRNACYDSPWYDWPFKIRKMINLIQTRASKDTKLTCIFFDLSMEIFLKVV HILYKMIWFPKTPNFIHQK >TbOr138 MEVLRFAKNLITNFTSNKKLDKSWLIFDNKSYILHKRVYMIAGFWFDQHFIVKSMIMFA HFMVSFSVGPSQTKYIVSHRNEFETLFTNIQFIVAHIYFVFLRIHRMFVGRKLKLLHNEIV VDWDKTQNEEERKILHTWALDTRRTTLNTMVMGVTFAMVVYCTSIMPKILDYLNPLDE PRPDNFLVNLEFITYPESMYFMMNCVYIDVGISLATVFVLTIFNTLILFSNHISSKCCIIMY KLERLNHLEIPHVKWNMKKCIEYHRRTIQYANDTINYFAPFIILTSLLVMILITLTLFDMQI NFDKLNKIAKLLPLLSLVLILMFVVTIYAERITTYTEALYTACYCTEWYKFSPTSKTALNII QVRCIKPTPVSAYGVVTFSMDLFVKIVQTSFSYFAVICTLQDQG >TbOr139 MYFFVLKSVIIHYLTKRITYESWRIFDSKYYRLQKRLYQFIGFWFQQPFIVKGFIITCHTVL SIIIWPTQMKYLISHIHDFDSICSNIQFVSLHVLLVYIRLHRLIARDKLKHLYHKISVDWETI RNPEELDILKYYSNMTRKSIFKSTANVTLLIIFYITPLMPKILDYFIPLDHPRPYNFLINLEF LLYGEPFIFIMNYVYNTLGSTFGIFYLAYCYNTLLLFTNHISANFSIIYKLENITRNDIYND VITVKLNKIIALHIDTLFVENIRTFFEYLLVIGTTDLIVSLTLTLFDIQINIDKPYIVAKLLGE LLTIIAIATTVVFYAENIIQFSSLIKQSFYLAWYEWATKSKLTLRFVQLRTVANFFFVPNIY VVFPIQRLIG >TbOr140 MRISFIKKLNHKTKKETVRKKIDLFRTKYLLYSVTVYIMAVWPGQNKLVRTLLFCGQIFL SCVIITGQYGHLFCKVETVNEGLFDVLYIFLHVFTQICRLDFDTKRKLLKKLHTQLEEDW DNISSDTELKIMKRFAVIASHTIRKTATFIVICLIIVILTSIYPKILNAIDPLEKPRPEIYIFNLY FVWHPTASFFKKYFYNFLASVVGALFFLFLFSLYLVMYNHIYARFFVIKSILEKMYDFISP QFVNEIARDFRTSIQIHIKTTAFCSMTLKYFTLSLIFGIICDMVGFALALYTIQVNSKTLSE VLKTLPIFVCLLITIFILCMFGERLVTLSSGIRYDVYKLPWYCWPRRPLTVFNIMQNRCNT PCEINPMGLFSCKMEFYVTIIQTSFCYFTALIAVQETSNK >TbOr141

446 MKNFLMKKLKLHRKKKGKNFDVFRTNYFIYSLTLYIMGAWPDQHQLFKTLLFSGQIFLS ITIMTSQFGYLWFQVETVNEGLFDVLYIFLHVLVQIARLDLDIKSKLLKKIHEEIREDWKN ISSDTEKEIMIKFANYGTETIKRIAVTLFSCVFMVLFTAIFPKILDALDPLDKPRPENYVINL QFILHPRASFFKKYVYNVIAIVVGGVFFLFLFSLFIVTYYHVYARFFFMKSILVEMSDFTS PMFANKIVQNFRTCIQCHVKATTFSSTTLEFITLSLVAGVVCDMVGFALALYTIQLNTKR LAVVFKTFPVFLVLLLVFFFLCRIGERLVTISSSIKYDMYKLPWYCWPRRPLTIFNIMQNR CNTPCEINPMGLFNCKMEFYVTVIQTSFCYFTALITMQDTGNE >TbOr142 MLKNFRTKLWNLLFVGTEKPNSLFKLDYFTEYKHVLALIGYWPYQTYHQFLIIKFMQIV GSLTLAKGQIQYLILNRREFDAMCSNLDFVVLHVTLYFLEPNVISNKRRIKNLFEEINIYW QSVTKEEEKRIFHRHNVIFKKRAKQMEVSFILCIITLLICPFIPKLLDILYPLEENRPIEFLVP VMLKVLPSTFLVFYIINICACTGGIAYFIAVSHILHISNFYCTKIDLALFSLSDNKSELTKKG MNPGVTCNQLMLTIRRHREANMFFERVMDVVEYMVFIFAIAATVGWCFGFYLVMLNL TTMKNYSIMPMFAFSYYLMNVITLSGERIIMYSEKFRSSCFRSEWFTLSTPSKKNFNMLL HNCSNYRHISCFRIFTLSFESILKVFQTAFSYYTLFWSFR >TbOr143 MILLRFFFNILTFFNIPRNYKIYNDKSLVLNTKYYAYYKLSLLLPGYWPYQSVSQIFISKA FLGIVSAFAGKCQIYYLMLHKNEFDVICSNVVFIVTHFFAHIAEINRFFNTKTLKRLHDEI NIDWENSDFTKTRDVLMERTKRHCDVIKKRTVFCYSVVFVLSFSPLITKILDIIKPLDETR SINFAIPFEFEHLPNTFLTLFILNLAALLAGVSVPTAIINIISLSSYCSTKMAFIEQKLQMIES EINCEKNEQRKIKDLRMCAVLHNQFIQFFETFQKYISICIFIAALLDVVCLSSTLYMMQVS ELSNFAKFLPLVGVTMFYCFIICSSGEEIKSSSNDLNYRCYTTIWYIWPEKSKKMLHMIQG KTAMEYQMKGGGILALNSESFLVLIQSAFSYFTIFWSIRNSS >TbOr144 MIKKVNARLIFESEYFKLYKLVYKIIGCWPYQHFCGQIFQLVFFGICSMAILLAQIKYFYH NRNDFDEIATNFHFIIIHILLHCLRYHRAIKGKQLEQLHDGIANDWQMNRETKYYSLLHD YAIKFKKIAYRITVGSGIVALSMYFKLFWPRLCNLIMPAKEKRPDTLIFNLRFTIIRSKNFP FALNMIGIFTGILFFVVFIMTFLILVYHTCCLFTLVGAQLIDIDNRDHNKCQKQIAACIKQH YISLKFAAAVRSYFKIIFLIALSVLVAVNTLLLYMLQFKLNNEKERALLDVVPTLAMVLF FTYTPCSLGEKLMEKSFDVVKQCYDSKWLYWSNINKKSLHIIQSRCYLLCLINAGVFTL KNEFFVDVLQTSFTYFTVFSSTRNKNN >TbOr145 MKSLIMTVSNKSRTANTIVDNSEKDSDIFGSHYYKIYKDFYKIIGCWPYQNLLGQQALM LIFMGFLTAITFPAQVNYLYSYRQNFDELLVNVQFLLMHIAMHFLRYHRLLNGKKLRIC HDQIKNDWRDFKQHIDFEILHRYVLKNKHTIHLISAIANLLIFIVIVQVFWPYKKDEGTFIF KLEFHLLKSKFWHYFCNIMGAFVVIYMTMTHFMSILILTTHECALFTIVRSHILNIKSQDE NDKKRNLTECYCQFRDCIQLHTKTFKYAENVRSYFYFMIFYGVFIMSTVTSLSLFNFQYK FQHGVIFSSILPQLFTELYVTYTCCCLAEKLYEHHFEIKESCYESRWINWSTADRKILHIV QIRASNQVYILYAGPFEISLAFFVNMMQTSFTYFTVMSTLRKNDDEMNIL >TbOr146 SFYIVVIVQLFWPRENGEHLFVLKLEFHLLKSKFWPYYANICGPFVAITMVMAFSMSVLI LTSHECALFTIVGSRILNINGQNSYKTKSSKKYYNQFRDCIQLHTRLFEYAGITRSYFYFV MFYGIVFLVAVSSLSLYNFQYKFQRGMNFTDVLPTAAAELFLAYMCCFVAEELYVHHF KIKETCYKSSWINWSTSNKKMLNIIQIRSCNQVYIFYAGPFHLSLTFFVNMMQTSFTYFT VFSTVGKNN >TbOr147

447 MMSVSNKNCGANNIVDNLKIESDIFDSNYYKTYKYFYKIVGCWPYQNLFGQQLLGITV MTVVTTITFPAQINYLRTHRQNFDEVLVNLQFIGLHIMMHLFRYHRFLNEKKLRICHDQI KQDWKNLRQQNESKILHAYAQKNKEVLHMISAFSIFFLFFIATYPFWSNGKGEGKFMLK LEFHLLHSKFWHHFCNIFGGFYAIFLTMSFDMSLLILTTHECALFTIVGSRILNINGQNSY KTKSSKKYYNQFRDCIQLHTRLFEYAGITRSYFYFVMFYGIVFLVAVSSLSLYNFQYKFQ RGMNFTDVLPTAAAELFLAYMCCFVAEELYVHHFKIKETCYKSSWINWSTSNKKMLNII QIRSCNQVYIFYAGPFHLSLTFFVNMMQTSFTYFTVFSTVGKNN >TbOr148 MLLLNENVSFNKKIVKFGKFDIFDSHYYKIYKLVHKFNGSWPDQNIIEILIVFIYEAIVGIL TVAGQLEYLFNHRRNFNEVSTNLPFTIISLMVVFLRIHRILCGRKLKICHDAIRQHWNKA FNSEINKQILQKFALKSKNIIFSFTVIIYSVISIVNASIVWSLLVNMVFRSEHEREFYLIFQIE FRFINLYAFPYIANICGSYIVAIHFLVIMSTCIVLSLHSCALFALLGHQLLFTGNKILSSFSL VKKTSRYNQLREFFNTHLETYKFFNCMQSYFCLIFVSGLAMMVLVLSLGLYTFQIRMKQ NIILSESLIVICSMIGLALLIATLGEQMTHQGSEVRDSCYKSNWVDWTYSEKKSLLVIQQ MCNQGIIVTAGPVITISYEFFVKVLQTSFTYYTMFSTVGINH >TbOr149 MTEFSFKFEEREDREVIFFQIKMILSDKKVTKKLKFNDSQDSDIFHSHYYKLYKIAHQLN GSWPYQNIFKQLIVVAYEWIVGGITMFGQLEYLFSHRQNFYEFSSNVQFIFISLMMLFAR VHRLTNGKKLKICHDTMQKSWKNVNTESINQILHKFALKTKHMILYVTAIIYFTISFVDA RILWSQLANWNSGSKSENRTYLIFQTEFHLIKLSCASAISNILGSLIVAIQFLVFIISCIVLSS HSWALFTIVGYQLLSVGNETLTNYEIMKKEHHSKHIRNCTQRHLDSLEFADNIQAYFNL LFISGLVMVVSVLSLALYTIQIRIKKRLTFTEALVTGFLMIILAFLVGTLGERTRYHCSKIQ NFCYYSNWLNLPINDRKTLIIIQQKCHHETVFSAGPFILCNEFFVKVLQTSFTYYTMFSTM EEIQG >TbOr150 MTLSSENIRTKETFKYVKGSDIFNSHYYKLYKFAHKINGSWPYQNIINHIIVLIYEILVGIL TIVGQLEYLYSHRQDFDVLSGNIQYIFISLMMGCSRLHRIVNGKKLRSCHDTVKQNWDN INTKVFQYILHDYALKTKHFILHSTVIIYSTIILVDAGILWSLLVNTVLRPEDKREPYLIFQI EFRFIKLDGLQYVANIFGSLIVAIHFLIFMITCLVLSLHSCSLFAVVGYQILLIGSGTKRNSS HMEKVDCDNHMRKFVQRHWKSYEFFESIQRYFDLMFMSGLAMIVSVLSLALYTFHLRI KKNIIFSESLIVIILMIILAYSVCILGERMRYNCTELKLTCYKSNWMNWPIMNRKCLLLIQ QICDRETVFRAGPFILCNEFFVKVLQTSFTYYTMFSSVGELNK >TbOr151 MTASRKNKLQIKEIQHLRGSDIFDSHYFKIYKLAHKLNGSWCYQYIFVPILVVFYEVGVG SITVVGQIEYLYNHRQNFDDFSTNLQFSFISLITAIARLNRMINRKKFRICHDEIKQHWHES MNAEGTQQILHKSALKSKHAVLFITVFVVSTIFMVDLSILWSLFVNKVLQPEENRQSYLI FQIELKFIKFDSMPHIANIFGSLIVCGEFVNFITTCMILSSHSCALFAIVGYKLVSIGNKMIN YSFIKRSVCYTQIRECVVLHWKAFGFFDVMQSYFLVMFFAGSAMIVLVLSLSLYTFQSRI KRNIILSESLVVILLMILISLFVAALGERVSHHGSILRKSCYESNWIYWPNIDRKFILFIQKM CNQEIIFGAGPFALTNEFVIKLLQTSFGYYTMLTAVGEKSR >TbOr152 MTALNESKLQIRKTRPDIFDSHYYKVYKLAHKFNGSWCYQDILVPFLVVSYEVVVGSLT VAGQLEYLYNHRQDFDDFSTNLQFSFISIMIVILRLNRMFNRKKFRICHDEIKQQWDISM NEEGTKQILHESAVKSQYAVLYFIVFIFSIIFTVDISILWSLFLNKVLLPEEKRQSYLMFQIE FQFIKFDAMPHIANIFGSLIVCVEFLNFITTCIVFSLHSCAVFTIVGYQLVTMANTMMKNC SSMKQSICYNQLRECVQQHWKAFEFFDVMQSYFLILFFAGSAMIVSVLSLSLYTFQLRM

448 KRNIILSESLAVIFLMIMLGLLVATLGERTCHNCAEIKKCCYKSNWINWPIIDRKYLLIIQQ ICNQEIIFRAGPFALNNEFVINLIRTSFAYYTMFTTVGEKDG >TbOr153 MTLADENVTKNMKIEHLRMSNIFNSHYYKLYRLTHKINGTWPYQHIFQYLLVLGYEAL VGSLTIAGQLEYLYNHRQNFNEFSTNLQFTCISLMVIFVRLHRMFHGKQLRICHDVIKQH WDETFNADVFQQTLHKFALKTKHIVLYTTVCIYLTIILVDLGILWSLFLNTVLRTEENRQ SYLMFQIEFRFLELKAMPHIANVFGSLIVAMKLLIFSNTCVVLSLHSCALFAIVGRQLVSM GNTIMTNYSLIRETTCYNRMREFVQRHLESFEFFYSIQSYIWLLLFGGLAMVVLVLSLAL YTFQIRMKRNIFFSESLIVMLLMITLSLLVSSLGEIMSQYGSEVKYCCYKSNWINWSNIDR KSLLIIQQICNREVIFRAGPFLISYEFVVKVLQTSFTYYTMFSAVGETSE >TbOr154 MDLWEKNLLHQNEKEEESKKCDIFNSHYYKIYKFAHKLNGSWPYQNIFQQLIVVIYEVV VGSLTILGQFEYLYSHRQNFDEFSTNLQFTFISSMIVFARVHRMFHGSQLKICHNAIKQH WDETVNTKGFQQILHEFALESKHIILFSTIIIYSSIFMVDASILWSLFVKMILRPGENRESYL MFQIEYRFIKVDALPYIANILGSLIVAIQFLIFIITCIVLSLHSCALFTIVGRQLVIMSNKRVS NYCFIEKTASYNQMRGLIQRHLESFEFFDSIQSYFYFLLFSGLAMIVSVLSLALYTVRKR >TbOr155 MSGLKERSENIKKTRKSEDVTTKRHQDIYKSQYYALYRRICYGLGCWPYQKVYTRLPIF VIHSLIVCRSFWGQVNYLKLKNEDMDNFLKNLFFLIAEISISLLSCNFHLSKKPIRKLHDL LKKDWQTIETMQDYLLMRKHTERVQQKMRLLSVFSTPVVFLVIAVPFVPKILDRIKPLE RNRPDIFLFDVQVLRFKKPEWLMKYLTTMIAVLLGMLYLVMFTMLSVICCDHYRLKYIL IRARLENISRRYGLICCYVEKVSHHLKDIVNKHVKSIEYAYTTDECFGRSISVELLTFIITISI SIYILQLNMGKDMGAILSGLPGLYCSIGLFFFICHLSEILIQTSSSLYWHCYDIPWYNFNHN DVQITSLIVLRSKKYYSVGAFGLFTFSHEGFLTILQTAFTYCAMMFTVREKLS >TbOr156 MSMSSVKKSAYIEDTCVISKRVENIYDTDYYKFYKQIVTVIGCWPYQKRYQVYLIFFFLF VFSWRCFASQLYFVKCLTIDQMDKILSNVFYIFAHLIIMMIRFDCIFQRKKKRLLHDNLL HDWQNIQTIEDYLVLRKHAELAQSYILKLYYGLFSIIGIAFTSPFIGPVLDYVNPLNESRPD TFMFDTKFYLFKEPEWFMKFMTTSFGATLALLFLIMSFGYVNIFTIHYVAKHAIIKSRLHL ATKQFYSICYDKTVSSSSVQSVVPFHHEAISFGCLLTRTTEKLVLLCTVSSVLLISISFYSV QRNINDVRKMINHFPGVYCSLLLIFLMSHTGEIIIQTSTELFEF >TbOr157 MNFSFNPINIMLRTHSSDEHTIRPSSFDVARKRVKDIFKTDYYKYYRYVSTMHGCWPYQ GFYKYIIMLIHYFIGFRCFSGQVNHIISEPDIDSIIKNLYFLIAAITLLSLRLYSIFHRKEFLLI HDDVIRDWQKIQTIGEYLLLEKYARKSQTAIRIAFQILILMTVIVILLPFMPLLLDYMQPLE EKRPDAFLLEVKFLNFREPEWIMKYITITIGGIVSLMYLATAFGFAMTFSNQFAFKHALV KLKLDTISKRYRSTDFCNVENMMKLVVLVHKKAIDYSFLITISIKKITSASVIFGFLLIGTSI YSVQLNFNNSGKLLTSLPGTFCSLVSIFVVSHLAEKMGKSSAAVATYCSESQWYYFNNS DKQSLCLMMQRCSKIYKMEAYGFFTYNYEGYLTLLQTAFTYCTMMFTLKDKTI >TbOr158 MSCASSFDVKKAQIQHFDITSKKVENIFKSNYYKKYRYVSLIQGCWPYQGSYKYIITLIH TIIGFRCFASQVYYLRNHPDIDTVIQNVYFLIASVAMTTLRLNFFFQRKKILLLHDYLRHD WQKILTLGEYLLMRKFAEKSQKAIRIMLQSVLCVITFIFVSPLIPVILDYIKPLDEKRPDAF LFEVRFKNFKEPEWLMKYLTTTVGSTQGLLYLATFFGYTIIFVNQFGLKNAIIKLEFDTVS TRHRSAILCNNVANALRVVVITHKKTIEYSYLLQISINNIVAMSVIFGILLIGTSIYGLQLNL

449 NRPDRILNALPGVYCAFLSMFLVAYLSESLANSSAALKLYCYSSLWYNLSNNDKRNLCF MMQRCEKVFVVDACGIFTYSYEGFLTILQTGFTYCTMMFTFREKTN >TbOr159 MISMVILLAQVRYFLKNRKNLDETLTNCVFTLIHSYMYVIRIHRIFFTTELRELHDQIIFD WKNITDETERQIFCFYAMKVKKIINCIVVIIFGVILVLYMLMLLPTLLDLIQPLDEPRPLRFI VYFDFKYLKNHYNTLMFIYNIVGVTGAVFYFIEFQSSFFVLSFHSYAKFSLTIIKLQQLQQ NFNKTITTNNNTNLQESCYTNIKLHIKAILFAEKIDNYLKILSLVGLFMALLTLSLTLYVL NLKIGSVDMLNVTAILIGDLTLLYSFAAYSEEIMALSSNMGNHWYNTDWTTWSNPLKN LIVIVACRCMKPMKLTLCGLGSASFELFVTVLQTAFTYFTVMITFKENY >TbOr160 MNILYDKLFSYFFYYIKRRFNIERNTIFNTHFYKVYVWFFKIAGFWPYQTIWEDIFFLSLH IAISIFILPAQGRFLYNNRHNFDEMTANFVFAIVHIFIYVLRAHRFFYDLQIRQLHDEIIYD WKNLLNYDQKILYNYSMKSKKLVTYNTGFLITLNILLNSLPLVSLVQDLVYPLSEPRPYR FFLLFEFKVFAHYAFFMMFVYNLVGLTIGLLYLLVFTNTFVVLTSHVCAKFALITNRLKP FTKNDTVFNGNESVNELCHSCVKLHIQALLYAERTSIYFGILLLVGSLCSIMAMSLILYVL KLKFGTVSMFQLITLVFDVILIWLCCGVFGERVMYHSTCISDYWYVYKTDWINWPNSIK KSNIIISCRCIKQSRLSLCGLGYLSMELFVSMLQTSFTYFTVITSFTE >TbOr161 MVLKVKLIFGINYLTEDNLSAKNTNFESHYYKIYIFFYKIAGFWPYQKERQEIVTNIIEFMI SMFIITTQTRFLYRKCHNFDEIFVNLLFLIIRMYMYVIRYHRMFFQNQSRSLYDEIIFDWK KITHESDRKVLDIYATTTMRSVTFATGTIVFAFLPVFPKIQDLLIPIDESRPFRFIVKFDFRF LKNHFILMFYIYNFTTFVTGVLYFIVFYKNFIILTSHTCAKFALIIFRLKYYTNSGVVITNN DKYELSCDNIRLDVQAILYGERVNDCLEICTLIGLTFSLLFVSFIILYVFHVRFGTMGMIGL FFLGTMALMIFYNFTNYGGKVMEHSSSISSHWCESRDDIICILEIEIGHAIMRSSILFPCYPT YLPSIL >TbOr162 MKAINEVDEFYKTHFKIYRDIGTRCFAWIYQSRLIKIGSLLPTGFASLLTFNWQMVFMKN HYQNIDVVLRTCLFIFLHIFIIFSRLSHMINDNANRKLHHEVQNDWTNIECAEDAEVMKR CASQLKSHLQHFSCVKEFSIPEFMLTVTIIFAISPTIPKLLDSIKPLDEPRPTNFIVDIDFITDL QPEWLLKWIFSTTILIIGLPLIEMIISLVWFSLHHTVTKYSVLFRLETYKNYCQTHGTDFT MMKSIIRYHVLIFYNNLLKNSTEAIIFSGYLTILLMLSIGIYNLQLNFNNPGKLLGILPICILP IIVLYSVCLKSEDVTAISSNLPFYYDLPWYRFHRINIIHTFKFMQFKCGQIYIYGAYALVK ASLETVLSTLQAALSYCALFYSVREKF >TbOr163 MSRTEVTLNREPDIYKSDYFKFYRAVGRITFCWAYQKKIIRMASLLPIGFISMATLNWQL VFLKNHRHNLDSSMRAILFLIVHMFIIFVRLYFEFYDHKSKMLHDEVQNDWMNYDNVE GYEIMKRHALRLKAQMLIFCNLNFALVIISVITPTIPKLLDILKPLNKPRPPSFIIDIDFITNL QPEWLLKYIFSTMAMMVGQPMTQVIMCFFWFALHHSVTKYSVLSSKLDMVKNDNEM MMMMMMKSLVRYHVQTFKYNEIMKNNLETILFSGYLSLSLMLSLIIYNLQLNFSDPSK MLSLTPTLTLLLLVLFTVSWMCENIITSSSSLHSHCYNIPWYDFRNKNIIHSFTFIHFKCGH TYIFTSFSLIKYSLDVALTLLQTAFSYCALFYSVRDRI >TbOr164 MLRFIKKRLLDQDEDLALKQKIDHFYKCNYFKLYRSIARTTFCWVYQNKLVRLVSLLPT GVASIATYNWQIVYVKNHRHNLDSTLRAMIFLIVHTYVLCIRVYHEANDHVNKMLHDE VHNDWNNIESSKDQEVMQKYALRVKSQMWRVFKLNITTFIICLITPTIPKILDVIEPLDKP RPTNLVVDLDFISNLQPEWLLKWIFSTMILMIGLPLVQYIMCFSWFQLNHSLTKYCVLRS

450 RFETIKNKCRTQKSISAMMKSLVRYHASIFKFNKIFKSNLETITFFGYLSIMTIMSLTIYYL QQYFNEPSKMLSITPTLVFPIILLYTVCSFSEQIITKSTNLLFHSYDLPWYYFKDANMIHSY IFIHYKCGQNYVYTAYSLVRCSLELVLTTLQTAFSYCAILYSVRDEV >TbOr165 MLRFFKRKRINQEQELAWNQKVDNFYKCDYFKFYRSIARITHTWIYQKKILRIVSLAPIG FASFFIFNWQVVFLKNHFSNLDAVLRAILFIIIHTYIVFVRLYHEFNDHANKMLHDEVRN DWTNIETTEDHEVMQQYVLQLKTHMRMSFKLNVGLFILCLSTPVIPKILDYVNPMDEPR PYNFVVDIDFITDLQPEWLLKYIFTNTSLIMGLPLTQWIMCFAWFQLNHSCTKYSVLRAR FELIKNKCLKQESILKLMKSLVKYHASVFKFNDILKFNLEAFTFFGYLSLMMMLSLTIYN FQLYFNNPSKMLSLTPTCILPILILYSVCLSSENITTNSFNLHYNSYDLPWYYFKHTNMIHI FTIIHYKCGQTYVYTSFSLIKCCLELVLTALRAAFSYCALFYSVREKI >TbOr166 MNEYQEAIPKRELENIYHSHYFKLYKNVSRASFGWVYQNKLLRLASLLPTGFASMATFN WQIIFLKNHRHDLDIALRTVLFIIIHVTLIFMRLSNMLNDHSNKILHDEVQNDWKNIETVE DFKVMSRYALRLKSQMQTYFKLNIVVLLVCVTTPTIPKILDIIKPLNEPRPSNLIVDIDFIN DLQPEWLLKWIFSTTIILVALPMIEMLIWFLWFALNHSCTKYSVLRSRFDTIKNKYKLQN TISTMMKSLIRYHTMIFKYNEIVKSCLESIMFSGYLAVLMLLSLAIYNLQLNFNNPGKML SIFPTIIIPLIILFTVSLLSENVITNSSNLHFHSYDLPWYYFRQKKIIHTFTFIHYKCGQTYVFT SFSLVKYSSDLVISTLHAAFSYCALFYSVREKI >TbOr167 MFGFFIKKKNNEGKKVTPRCVLDDFYKCDYFNLYRSIARATFCWVYQHRLVRIASLAPA GIASVATFNWQLLFLKNHRDNLDLALRTILFLLIHAITIFVRLQIEFADHSNKMLHEEVRN DWINVETNEDHEVMKQYVLKLKTQMQMFFKLNVVLYILCFLTPTIPKVLDIVKPLDKPR PFNFVVDIDFFTDLQPEWLLKWIFSSIILTIGLPIIQLLICFLWFAMNHTCTKYCVMRTRLD MIKSEWKIGKKISKMMNSLIRYHAMIFKYICFCRYNEIVKHSLEAIVFLGYLSIVMLLSLA IYNLQLYFNSPSKMIGLTPTIILPLLILFTVCFLSENITTYSSDLDFSSYDLPWYCIQHRNIIH TFTFIHFKCGQTYIFTSYSLIKYTLDLALTTLQTACTYCALFYSFREKI >TbOr168 MFGFFIKKKNNEGKKVTPRCVLDDFYKCDYFNLYRSIARATFCWVYQHRLVRIASLAPA GIASVATFNWQLLFLKNHRDNLDLALRTILFLLIHAITIFVRLQIEFADHSNKMLHEEVRN DWTNAKTTEDHEVMKRYVLRLKTQIKMFFKLNVALYILCFLTPTIPKLLDIVKPLDEPRP YNLVVDIDFITNLQPEWLLKWIFTSIIVLIGQPIVQFTMIFLWFSFNHTYTKYCVMRTRLD TITNNYHTEQTILKLLKSVIRYHTSIFKFNKIVRNSLQGIMFFGYLTIMLMLSIAIYNLQLN FNKLNKM >TbOr169 MFGFFIKKKNNEGKKVTPRCVLDDFYKCDYFNLYRSIARATFCWVYQHRLVRIASLAPA GIASVATFNWQLLFLKNHRDNLDLALRTILFLLIHAITIFVRLQIEFADHSNKMLHEEVRN DWTNAKTTEDHEVMKRYVLRLKTQIKMFFLNVALYILCFLTPTIPKLLDIVKPLDEPRPY NLVVDIDFITNLQPEWLLKWIFTSIIVLIGQPIVQFTMIFLWFSFNHTYTKYCVMTRLDTIT NNYHTEQTILKLLKSVIRYHTSIFFNKIVRNSLQGIMFFGYLTIMLMLSIAIYNLQLNFNKL NKMLSLTPTFILPLIILFTIALSCGHIIQHSSNLLFHYNLPWYCFQHKNIIHTFTLIHFKCGQ AYVFTSYSLITYSVDLALRTIQTAFSYCALFYSVREKI >TbOr170 MICKKKKIDKNASYEYYYSYNKLFLSTIGLWPYQNDFLKFVFSFVTIVAGLSQVGVQIG GMVAAKISKDQKVFLESIAPVVVNGLVLPKFINFLFNKNKMNMLYRQVEEDWRTFKSS QDVQLLREFAIKTYKTCPVYAGAVYGGLLPVVLLPTIPAILEACGLIKQEEKLLPYPVQY

451 FVNLDRYYYHVIIDAFVGTFYLVMIVVAVDTMFVLFIHHACSVFSIIRLRLKNIVKELQV DSVAQKDDLSYRRLVNCIKLHNTMLRFVSCLESATSACFLIKIGFNVAGITFTGFQAVML LNKPGESFRYLTFTFAQLSHLYVQCSPSQRLFDHSLELHTDGYNGLWYHATNRTRKLLT MFLMRTLRPCKITAGKVFVLNIEYFSNVLRTCMSYFTVLSSSQSA >TbNmdar1 MHGSFLGIVALLLGVFVSSPVSCTAEANKTARAAKSRAKASWRVAVCLTQSQAPLKID VQKAWENARLESSKAELELSYVEARAPVTADSLSYPLSLLNKFCTDIENGRTVLSLIIGG GSASRFLVTAAASLNMPTLWIPNTHKDFLRQGKQRRFETRLGTSAREMGAAAAVLMHK ANWHAFTLLVDSTLLPIHYLLQSRVNLTPRTIIHLPTNEKTLRLRLRRIAEQGGSGNVLVL ACDLNNARKIITMANKFEMLSGRFLWLWLDVKAELRSSEPNLISSHVVHPARQPAVAPI DNVALKSLEVKAQLYPDSRPHLHSLPNLASDVHQLQDYWLKKEKSVKKREDREFHFLE EDEEDLKPDKTVNSKNFMPVGMLALRPASTRLAGGDTIMSKAVRELSLVLDDTIAEARP RLARFRDSQIKEYFVPECYASITARFLTNEARENVSKTVTKKLRRSMRQISKDRAEFHLL NLQGIVFPGNKTQLRWTKVGSVRGGKEVRLDTIIWPGGGIVPAYMEQGGERIGMPAYKI VTALAPPFTMSTNLQEGLCLRGIVCRIGSEVKCCYGYSMDLLTYISQELHFRFELYITKD GFFGKRSGRNWNGIVGELVSGRAQMAFAPISVSTKRAEVIDFTIPYYFSGVSFLTAPKLR SDIPLLAFLQPFSTELWIAVFTSLNITAMAVAVYEWLSPFGLNPWGRQRSKNFSIASALW VMWGLLCGHLVAFKAPKSWPNKFLINVWGGFSVIFVASYTANIAALIAGLFFHSSINNY DHRSLLSQKVGAPRASIAEYYIQRGNPQLWSHMVRYSVSSVADGVERLRNGSLDILIAD TPILDYYRATDNGCRLHKIGDQISDDTYAVALTKGHPLKESISKLIANYTSTGLLDILQNK WYGDLPCLRGGVETGIEHDHGGQPRPLGVASVAGVFCLLMLGVFLGTIILAGEHLFYKY TLPRLRHRPKTSIWRSRNVMFFSQKLYRFINCVELVSPHHAARELVHTVRQGQITSLFQK SVKRKEHEQRRRRKSKAQFFEMIQEIRR >TbNmdar2 MNYEQYGRSTASAQYFLQLAGYLGIPVIAWNADNSGLERRASQSSLQLQLAPTLEHQTA AMLSILERYKWHQFSVVTSQIAGHDDFIQAVRERISDMQDKFKFTILKAVLVTKPLDLLE LANSESRVMLLYSTKEEAPTILQAARDYKITGENYVWVVTQSVIANLQTSNMFPVGML GVHFDTSSNQLMNEIATAIKVYAYGVEEFLNDSENPNNSLNTHLTCDGVGESRWNTGD KFFKYLRNVSVENSEQGKPNTEFTQDGVLKAAELKIMNLRPGNGKQLVWEEIGVWKS WEKDGLDIKDIVWPGNMHTPPQGVPEKFYLKIGFLEEPPYINLSPPDPVTGKCLMDRGV HCRIGKESDMAEMDIQSAQRNESFYQCCSGFSVDLLQKFSEEIGFTYELVRVEDGKWGV LENGKWNGLIADLVNRKVDMVMTSLMINSERESVIDFTVPFMETGIAIVVAKRTGIISPT AFLEPFDTASWMMVGLVAIHLATFVIFAFEWFSPSGFDVKVNPSGGHQKMTNTSQNNR FSLCRIYWLIWAVLFQAAVHVDSPRGFSARFMTNVWAMFAVVFLAIYTANLAAFMITR EEFHEFSGIDDSRLSKPYSHKPMFKFGTVPWTNTDSTLSKYFKEMYSYMKHFNKTTVAE GVEAVINGELDAFIYDGTVLDYLVAQDEDCRLLTVGSWYARTGYGLAFPRNSKYLQMF NKRLLEYRNNGDLERLRRYWMTGTCKPDKEGQKTSDPLALEQFLSAFLMLMFGILVAA LLLFMEHLYFKYVRHHLAKSDKGGCFALLSRSMGKSLTFRGAVFEAQDVLRHHKCRDP ICDTHLWKVKHELDMAHLRVRQLEKDLEAHGIKPSKTKKTGNLGWWRGCFQSPKHHP TLEPLEVMAPHPLPPCFDSFLDTVEVSRPRDMTRTRIACTEFSETFLPTEIYRISEDTSVRT EIAEIETVL >TbIr1 MKGDLGCLLAFVFVLQSVQYTLTVDALPPIVKIGAIFTHDQRNSSTELAFKYAVYKINKE KIILPKTTLVYDIQYVPRNDSFHASKKACQQVKFGVHAVFGPSDPILGQHIHSICDALDIP HLEARLDLDTEAREFSINLHPAQSLLNKAYQDVMAFLNWTKVAIIYEDDYGIVKLRELV RSRKTEDIEVYLRKADPESYKQVLTEIKNKEIWNFIIDTKTENMHLFLNMILQLEMNDYK

452 YHYLFTTFDIETFDLEDFKYNFVNITAFRLIDAGDLGVRSILRDMERFQPAGNTILNKSRII EAEPALMYDSVQVFAVGLRTLEQSHALRPANVSCELEKPWDGGLSLINYINSVEMKGIS GPIEFKEGRRIQFKLDLLKLKQHSLVKVGEWHPGAGINVTDESAFYDPGTSNVSLLVITIL ETPYMMMHHEKNYTGNSRFYGFCVDLLEAVARKVGFTYHLKLVPDEKYGAMDPETGE WNGIVRELMQHEQPYVMLRSQGNYSGNARYEGFCIDLLKEIAHKVGFTYRIDLVPDGK YGVYDYETGEWNGIVRQLMDKKADLAVGSMTINYARESVIDFTKPFMNLGIAILFKVPT SHPARLFSFMNPLAIEIWLYVLAAYVLVSVTMFVVARFSPYEWNNPHLCHPGPEIVHNQ FSLSNSFWFTIGTLMQQGSDLNPKAASTRIVSGIWWFFTLIIISSYTANLAAFLTVERMITP IENAEDLASQTDISYGTLDSGSTMTFFRDSMIETYKKMWRFMENKKVFVGTYEEGIQRV LEGDYAFLMESTMLDYIVQRDCNLTQIGGLLDNKGYGIATPMGSPWRDKISLAILELQE KGEIQMLYDKWWKSPRDTCMRNDKGKENKANALGVDNIGGVFVVLLCGLAFAVLIAI FEFCYNSKKHTLSERQEQTSLPPPTFSGSLQGIHQIVQQQQQQPNQQESLCSEMARELCR ALRCRASSRRRQACEKCSTHVVGYPADSVTTPINGMRSQRTNAGGSVELSPHLHLHHHP PPHDFDGN >TbIr2 MVSRGWSIFLLCFGTLSSVGAFSRRVQIGGLFDGNEQLQKVFEFSTKYVNKHRHELDLG NKVHLVAKSIETTTDAYEVAEKVCELFSSGVAGVFGPSDPVTARHVQSMCDTLEVPHIA VSWDPEQERLKAVNLFPQADSFSNVIYQIVKEYEWRTVTIIYDTSDSLIRLHRILKMWDP KGYPITVRRIGPDPNYRPILKKVKDSGEENIIIDCSIDILAEVLKQALQVGIISDHHRLIVTN MDMQTLDLEPYSYTGMNFTGVRMIDPENEKVQELYAGVADLGLSAANELRVEHALMF DAVHLFAQAFKQLSEAIVGEVKQLACNSSTNWEHGISLSNFMRSTEMIGLTGLIKFDTAG YRSDFELEIINVGIEGVQKIGTYNTKSGIEWTPKPTSQIPESQLSLRNTTFRVLIALTKPYS MLKESSKKMTGNEQYEGYAIDIIDEISKMLGFNYTFMVQEDNVYGSLQTNGQWNGMIR RIMDNEADLGITDLTITSDRESAVDFTTPFMNLGIVILYKKPTKAPPSFMSFLSPFSVSVYI YLLLTYIFISLSFLVIGRMSPAEWNNPYPCIEEPENLENQWDLKNSFWFTIGSIMQQGSDV APLGPGTRVLAASWYFFCLIIVSSYTANLAAFLTVETVVRPIKDVDDLVNQNQIKYGAK NGGSTFQFFKNSNDSRFQKMYRYMMDNAKDVLTDSNDEGKNKVLNEDYAFLMESSSI EYLQERECNLSSVGKNLDSKGYGIAMRKNSTYRNELSAAVLKLQENGILTKLKTKWWK EKRGGGRCNDESGGGSATELTLANVGGVFLVLVIGVAVAGVATIIQFFLKVIYSAYRDK EPIKDALIEEMRFAMKCSLTKPVKKRKGSSNESGKSKSGSKNSKESESTPPYGVPTVITPD DDKKK >TbIr3 MTDYHSYLITSLDLHTVNLDEFRHGGTNITAFRLVDPNVTQNIVQDWIYGEQRYQRKLD LGQYNNKTETALMYDAVHLFAKALHVLDASQQIEIKSLSCDSSDTWPHGDSLINYMKV VEMAGLSGLIKFDHQGFRTDFDLGIIELNTKEGLINIGTWNSIRGINFTRSDRDVLETIVES LQNKTLRVTTILSAPYCMLRESSEKLIGNARYEGYNVDLIHEISRILGFNYSFNLVTDGKY GSLDRKTKEWDGMIKELLEQRADLAIADLTITYDREQAVDFTMPFMNLISILFRKPIKQP PNLFSFLSPLSLDVWIYMATAYLGVSILLFILARRVSAYYIFIYHLFVFLFTPYEWCNPHPC DKNPDHLVNRFKLLNCLWFTIGSLMRQGSNILPFSPYEWENPHPCNGQSGMLTNEFTLM NSLWFTIGSLMQQGSDIAPAVSTRMVAGMWWFFTLIMISSYTANLAAFLTVERMDSPIE SAEDLAKQTKIKYGALKGGSTAAFFRDSNFSTYQRMWSFMESQPISVFTESNIEGVDRVI KAKGTYAFLMESTSIEYVIERNCELTQVGGLLDSKGYGIAMPQSPYRTAINGAILKLQEE GKLHILKTRWWKERHGGGSCRDESSKSSSTANELGLANVGGVFVVLVGGMGVACVIA VCEFVWKSRKIAVEERKQKFSGKPICDGFTDLH >TbIr4

453 MMAKKWTIAFFLSIYLSYTQSMSRTIKIAIFSRVDTEHHRAFLDALEQVKSDNTAPAFDL EPIIRFIDIDTDSYNTAVACDLISEGVAAIFGPENKYTRGIVSSIASRFEIPHIEYGFRRIEDSE LPPTSINVYPDSKIYQQAIADMTKMMNWNSFTVIYDTADALSRIQDALSLHDPSTANVV TVRQLGEGPDYRPLLKEVANSLDYNVLLDISSELLPEVLRQAREVKLLNDYYMYIITYLD ATLLPIIHELNGTAANITGFKLTTNNDNRYTQVESSLLYDAVNLFSFTLEVLHKRIKGEKN FRLDPVPLICLGESKYTKYPAGQRIVDSMKELSKSGLLTGVMTFDEYGRRNFEPRVGEF QSNKFIETAVWNQEKGIHSTRTEEEREIELRENIERKVFKVTTRVGAPYVILVKDGSNRG VLIGDERYEGYCIDLIDAVAKELNFKYEFELVPDTNYGSYNEDTKSWNGLIKRLLDRKA DLAICDLTITYARQKAVDFTMPFMNLISILFSKPENKQPDLFSFLSPLSTDVWLYMMTAY LSVSVSVYFLAIAPSEWTNPHPCDEEPEELENNFDIMNSMWLTAGSLMQQGGDILPASSI RMIAGIWWFFVSSYTANLAAFLTAVKMDVSINDVNDLAKQTKIKYGAVKGGSTLAFFK NSNTSVYQRMYAAMLDAKPSVFTENNDEGVERVIKGKRQYAFIMESTSIEYNIERHCEL QQIGDLLDSKSYGIALPFSPYRTMISKTILKLQEKGMLEQLKSKWQEMHGGGTCGKEEP VQESAELGMANVGGVYVVLMFGFFAAILIGIGEFLWNCHVVAVDEKVSYCEALVHELK FTFSFHIKTKPVKITKSSNSNRSGSSGTSGFEGATPSRKSLVGSAVGSYLRLNILDKFSKEN ETNSRNNGNGNGNKLS >TbIr5 MNSTNSSVVSIPHYLIATHTLDDPYVMFDNVTRTYYGYSIDILHKFKQHFDFTYEIYAVN DTNVYDVLYKDILDRKADIVLHNSYIVHPLEDMIRFTKPVYEATGISMMTYRPGYSLTH FKFLGTFETNVWLSILLLYFVTSILLWFFDKWSPYSYRNNKANIYGDKRDFTFIESIWFC MINLTPQGRGELPENLSGRFIAMVWWLFGNCVLTFYAAKLAQFLTIPTSTMNITTIEDLC NQDNIQYGVTMNSSSYRYFKQTAAVNKMFHDTLIKMIYDDNLTMVERMTFATWTYPH KNKFTRLFHSMEAFGHVESFNAGIDRLQNTNVRYVFISETSKIKYFISNRCEFKVVGEEF APFQMRIVVREKSPLINKIDSVLAHMFRLNELQNISDKWLNSDFCSRNNVNDGNEALSV KLQNIIGLFIITFLGAISAYAVLIVEIYLHRKKLKSLLDSGPRRLNHLKASTDASLEYLNSF >TbIr6 MKHNTLIIFLIFFNGIIVFTQHLINETHQLVNVYINSLDNPLASKSVENSLNYIGAEYPNYL GQIYKVEINSLEPQNSVKEICSEWNRSMENAPHVIIDTTLTGKNSKFVQMISTKFGIPTILT RFRETNFVKNKINTSLDDENPNIISIIPPTNLLPEVVKQVVLNLDIDVAVLIDSDSYFDRIY GNMFLDVPIRKIIMKIKPVIKDLNEQIKNLWRLEMNNKRQKSAWFAVTLDECQLQCYNC MNFSIITVNPDIMEQSQQSLYVDLINKGVLSKPYLLSAFYFEVTKLAIFAVKEALKLNLW DSTVNQINCYSYYSKEKEQKLSQNFDFFGLLQNVMRTENTNSLFDNYVWIEPEGFYPKF HMTINLLSLLLMLLLYQKQKLDLGMHNLSGKLSENHSIKNFYRIVVKDDPPFVYYDEDA KQFSGYCIDILNELQKEPKFEYDLYQLSLPILQILQQMTIRQADTGIQSLDDLRKQFEYKY SVTVNDTTHYHFKKMVYIESKFHAWRNLLLDDKMDEYEQSKYLLWNYPLGKEFSDLV DGFKSAGYFDTHKSALESILNRSNKLAVIDNAARNEYYASLNCDLKVIGKEFTRRPFAIA VQDGSPLRNQINDLLKLTNDKVLENLKLKWWKQKEVCGTEVGINIEDIGGVFIIFFVLLG IFNVFFVSEMFYNREQKKFQHK >TbIr7 MSTKTTFYSAIFVLFLTTVFSQRNEDTPANLLIVVEENDQFILNLVNEVVSFAEHEIRDRRI TVNVKQLVISRDYPAGNIEAACAALSRGVHLILDFTWTGWIKLKKLAKEWGIAYVRGEP VIYPYVQTMDEILLIKNATDCALIFEYERELNQTLYYLIGNSIIRLVVIDEFNSMTVAKVG EMRPLPSYYVIYASTEKMEEFFKTAIDGNLVNRDGVWNLVFTDFNYKRFSYIANRIDFNI SVTILTMKNSVCCNLLHKKDCDCNKDFSIFPAYFSNLVTFLTGILNDIEKDKGLPAPISVP CNYINSSDSNPNNVTNSLSRFSEDITERINTYNNNTLAYYEDNLMLSYHAEIEISTFQSGE LDVLGSWITGKHIVMDPDKKIVPAKRYFRVGTAEAVPWTTKKLDSKGEVMKDVNGNII

454 WEGYCIELIQRLSELMDFDYDLVIPQDGSSGIKLPSGEWTGLVGDLAKGETDIIVGALTM TSEREEAIDFVAPYFEQSGILITIRKPVLKTSLFKFMTVLRLEVWLSIVGALTCTGFMIWIL DKYSPYSARNNKELYPYPCREFTLKESFWFALTSFTPQGGGEAPKALSSRTLVAAYWLF VVLMLATFTANLAAFLTVERMQSPVQSLEELARQSRINYTVLNGSSIHQYFINMQKAED KLYEVWKEITLNSSSDHVEFRVWDYPIKEQYGHILQAIEQAGPVMSVEEGFKKVVDSEN AEFAFIHDSSEIKYAVTKNCNLTEIGETFAEQPYAIAVQQGSHLQEEISRKILDLQKDRYF ERLSMKYWNQTLRHICPNSDENEGITLESLGGVFIATLIGLALAMLTLAGEVIYYKRKNA TGTENKEKMNEKIAAEMENDMMIFKKLTSQIKLKPAPVVPFDEKPFSKKNVQVSHISVY PRPFPFQQ >TbIr8 MADRETAVDFTLPIDDFIGFRIMMGKPKNLRNFFEFITILDYKIWLCIMASYFSTCLLLW MFDMWSPFTYRNNRCDYDIDKKIFTFKECLWFCLMSLTPQGCGKVPRNLSGKLIAATW WVFAFIILSTYNANLAVFLTEAGSEVTISSIDSLINQQKIQYGVVANSVSYHYFENLARIE QKFYDIWVEMTLNESISSSERMQYAVWDYPYKNKYSTMVKTMKSYGLPSNLDEALCKI RTSKVPYALIESAATVKYLTLSKCDYIIVGNEFNERPVTLIVQKGSTLTKNLNNIIHKLSFY GELQRLHRKWWDNITIECKVKNLSLDPIDFHTIGGAFSFLLIGVTLSYFGLFTEFFHAKKR ININE >TbIr9 MSIESDIPHYRIATALAPPFVFYDNLTDTYFGYCIDLLHEIQSEMNFTYEMYQVEENLYG QMDNDGNWNGLVRDLLDDRADIAVSSISVMAEREFVIDYTVSIYDSVGFSILMRKPKQP GSFFKFVTVLDGDVWLGVIAAYVCTSVLLWALDKWSPYSYRNDEGPDDAEKRVFTLKE SFWFCLASLTPQGCGGTPKNLSGRVVAATWWLFGFIILASYTANLAAFLTVSRLHAQITS LDELSNQHTIQYSVRANSAALQYFERLANIERIFYETWKNMTLVDDLSDLDRSRLSVWD YPLSDKYTRILDAIKYYQPPGDYDESIDRVLNEETPFALIEDAALIRHTALTNCDLTVVGK EFSMKPYAMMMQKNSPLKEPFNKAIISLTFRRKLEQLKEKWWNDDDKCDEDDDLTGEI KIESIGGSFLVIFIGIMLAYFVLLIEYCFFHDREGYQ >TbIr10 MPISNVKIFILLVLGVNLQPSDKTFSFAQKTPNFSSKKVNVFFIYDGNNEMVNRSVRNSL NTMKDRYPGYFGKVTAVEIDSSNPRKAADQICSKWFESLVRGTDNVPDIILDTTMKSKS AEIVQMITASLGMPTISTQYGQEGDLKNWKNLNRDQKKFLIQINPPPDLVPSVIRQLATQ MNISNAAILFDRSFIMDHKYKSLLLNVPTRHVIVKIDPSSVKSQLRKLRELDIVNFFILGSQ KTISSALEAAQNLTFIGRKFAWYTISLEVDLPQCESCTNVSLLSFTPHLNERQHTEPMKKL IQDGVLTKPYLVSSFYYEIATLAVVAMKSAMESNLWDPTKRRITCDTPESTRPPNVPND KNVVKDFDLLSILQNVTESDKFASLFDSYNWGKGNGYYYPTFEMDLKLLEFSQGTTTPD LVGNWTAGTNNPLMVLDESLIFNHTALTTYRIVTIINPPFVMYDNETKKYRGYCIDLLEK LQKEMKFIYELYLVEDNLFGTMDENGNWNGMIRDVKDKKADMALASLSVMAEREAVI DFTVPYYDLVGLSILISKPQSTTSLFKFLTVLEDEVWYSILGAYFCTSILMWSFDKWSPYS YQNNRDKYKDDDEKRLFTLKECLWFCMTSLTPQGGGEAPKNLSGRLIAATWWLFGFIII ASYTANLAAFLTVSRLDTPISSLDDLSKQYKIQYSIAENSTGYQYFERMANIEEKFYEIWR EMSLNDNLSNLERSRLAVWDYPVNDKYTKMFQSMKDAGFTKSIDEGIERVRNNSIPFAF IEDATRLKYITLTSCDLTMVGEEFSRKPYAIAVQQGSPLKDQLNNAILTLLNHRELEKLK DKWWNIHRNCEKEDDQSAGISIRNIGGVFIVIFVGIGLACVALVVEYYYYRDRSRVQKIE RKKSDRESINRGQAASNRASVLFEINQMRRF >TbIr11 MILVVLLQLVLTVTDVNVLAIFINQTATIAFHVDSEFFENDLDHEFYVTEVPRAIQELVQ NYMKFTALDIKPLKNLKANLRKDYLILITIMDCQKTWSTLSKAKREELIYIAVTNTPCPRI

455 PAEDGFSIPFTNEKSEISQIVFDLRSNDKRNWRKINIIYENSTDSQFVEELSELFLKNKYHE DDDISTTLFAIENKRNQTLVKEELDLILKRYGTEESGHRFVTIMSHELVPIVQEMAKLKK LMTPEDEWLFIIESNKNPFNISKFIERIGEGENLCFVYKISAPTQNCEENVLCQIKQVIKILA TNIDAQSARFSNQDYETKSQHLKSVKLKNKISLTHSIKQYFKKNNNRTCSKCYKWNMR SSLSWGISYTPKTNNTFLDVGSWTLELGVTLWDSLFPNRKYGFRRSNLPIATFHNPPWQI VDIASVRNRKYSGLVFDILNHLAKKLNFTVDVQFAVTDETKHTMKLLHQNNLSQMVKL DEAAMSTAQMIPENIISLVVEKKVFLGACASSTIKLFKKGVNFTLPISIQSYSLLAPNPVLL SRAMLFIAPFTSETWVTLLGSIFVVAPVLYIIQKADFDSDIFPANERGFHTLWQSIWYIYG ALLQQGGSNLPVTSSARLIVGMWWLVVMVIVAMYSGTLVAFLTFPRMDEAVSSISDLL YRKDEFSWGYPNGSIVEDYMLSLGGDKYTEFVKLAERHNTTEVLSIIKKIKGKKHVLLD WTSSLRFLMRMDYLITGQCYFTLSKAEVLNEPIALLVEPSNPYIKIINDELERLHESGLIKK WTSHWMPSKDQCWDGNVINLEAKKHKVNMSDMQGIFFVLAIGFTFAINFICCELLIHKK KTKKSLLLIKPFIN >TbIr12 MNVQTLHEIQKNALPSHIVVTTQENYPYSGVKLVNDNLVGTGYAFIILDLLAKYFNFTY TIKLPDKDTFEEVVGLLQNREVDVAVAFMPLFPELRMNCTYGTVLDESELTFFMRRPPD SASGSGLFAPFDISVWILIIITVFIVGPLIYICVTLRAKFLKDNPSEKYSFNTCMWFVYGAL LKQGSTVTPVSHSIRLLFATWWVFNIILTAFYTANLTAFLTLSQFTLPINSIKSLVEKKMN WFTFEEKAIDYMIRKNYAITADLKDSAYGSRIKYTKDIVNKNILNKIDESNAVFITENVY AQINLYKYYIGKMSKDLKEEERCKYVIMPNHIFKISRAFAFANKSSLSRQFNKLMRKLVE GGLIKYLKSKDLPEATICPLNLGSKERNLRLMDLSVTYSIVLGGLAIAIAIFIGELIYVRRC SKKVTPKDVAKRAPDAKTPKPQPLYGNWITPHAKTNDNANNDDFIPTIKVSPVSPMNAK IKNVNGREYYVVEVSKNNVRLIPIRNTSAIMFQHYRDLKK >TbIr13 MLIKQIIFWFLLSYNVTNQQLIRPVNVYTSIIESIHLIFNNTCIIFMHTVDEPIKAQELRESEE LKILQAHFSGLYIRTALMNLKTFEKKVGPAYFHIKRPLFVLLNDGDPMKYQFSRGIAKWI TMAYPTWVLFLRNSTSVDDFFADIFIPFDCIFLIVKQSEDTLNELIYEVYQINKSKEKITAQ FGTWNEQTGLEFTKLSLYQRRSDLFGHEIRVIAIQDPPLSIIERNKNGQMIDLKGFFGTILR IIQRSMNCSYSFQEASVWGTISSNGTTTGIIETLLNNKADLAASALTMTTNRLGQIAFTTP LYTARCRTYIRKPLYTAVKWEAYFAPFDFGIWESLAILVVISSFAILAARSLATILDDELK NLDQSTSTLSETMLLVYGAFCSQGMEHTIIDSVRIIHVMIHITGVIILSAYSAALISFLAVK VFVLPFTTMEGLLKDGTYRFSVVGESSDYELFQNTTDDTLKNMFEKALDVQEPLPNNY YDGLKQVCEKSKYAFMTLDNIVAVLKSKINCTLEPLDVIMQTTNGMAVRLRSPYQGIIN SNMLMMKDSGVMQRLFANEWSKELDSRKSAWSPVEINDILPLVSLLAGGIIFGGIILCSE YIFRKFTKNCHKKISRIKKNIDVKRNGLTVD >TbIr14 MEVYLQRGLFIIFISATLSFDNDHFLKLVLQSHRHYKTQSIVIITSEKSDIDSEHQLKHLIQS FSKGHIFSTVLNLEEFEMDVHLEMKMKKFIHPLVILFLPFKNQVKQFAVGNFQSMNPVW MMILPSGEVCMENKLGINLHLRFDSETIVYCPGEEKIYEWYSLTGEDLQINQLGIFDAHK NEIQFSNKTLYQRRNSLHGLEFNLIFYPGAYNYLRHVEKRKFDGYFARSFETLLQFANLT IKSQISDDDLGQYDPARNTWSGIFAEVHKSDFNIIISPLTIISKRAKVIDYVVPIGQSKMAL VFARPTNSLSWNIYFELLEYKVWIVTTIIILCMIISLSLVEYKFLNESFNLETSSHRFLTLSS LFCMRAHSDKLKSTSMRILFFSTFILALVIHALYSAFLIKHLRTIETFLPFDDIDGFVKDGS YRLITLNNTVYHQVFLDHEVRRSKGYEKMWKLANDPKTFPNDAYTGFSMVCKEKLAF FIPSIYIHFYKEQLPCSLSMIYTGYVESYSFVLKKGNPFKDFINLAIAKVFRHKKFVQHNSE

456 MEKLKVQACGKPVCEVEISKVTPVLLLLGIGISFSLFVLLTELSYFRYKLTKHKTNMTRM CNKCNNINK >TbIr15 MHSQFFRIRLWMQLVLLSLLLVFIINVLKYKYERTHWNLVRTIVDTYFDVFTMLCVQGC PDDSSNRCSIRMIYTSVMITALLLHSIYSAVLLKQLMSFEVRLPINDIDEMLRDGSYKWIT INQSGFHLLMKSVKSENLKSLWKFAANNNSFPQVVDEGMRIICNEKMVIFLSEMQVSLY EKDLPCKLASFYTGHTLSYSFLLTKNNPYTGFTSYAARSSFNHRKYTIDKKGIAIRGIKVC ENNLCPIELAQMTPGLVIQTMGIILSLCILLVEIIYCICKVKYKKLKKKITTSRSRLKF >TbIr16 MKVTPSNYLFIIIISSHGLHCSPCLMTLQFILDYAIERSWDQLVLFDVLDSECVLKYMRPF MQCLAENGIKSTFQPAINPNVPGVFNINRHRVSAIVFLDNFNLTAPDNVLNLASQKLLFN YYISWLFVNTNVSVQTTNTIMKELAIAIDSDVILATTFLDNYELSEWSVSYRDFCDSLKR YILKREVSRFENAHLDKGGSVDFVSVQNHTAFHLSHIYKIRINDNSSLIIHHLGGWNPFAS RLMRPIGVELRKHFQGFPLKIGLYNGDLEARQDQTVNKQLYDNVKPMLKYIEDIAQMM NASELTQHDKLGSLTNKAWTDLLHDVVNNNVDIGVGYITTTEERQEYMSFSFPVVKYM RNIYFVPPTSGKTRNIFLQPFNSELLICVCVTYFFIVATMEMITFVAKITHHHEDEKHAGL GEATLWCVSILCMQSPWTPISRSGKTVLLVGLLFALVIYNSYSGFITSILSVQTHAIKTIDD LKQFKFKIGFDSDDEEFIRTTNDSNLRDIYINAYNSRESGLNAIDGLLKVLNGSYGFFVSA SLARHFLKTTLIQEQCLMRELKLAQTESQVAFPMSKNCPYQKIINFILRTREFGILSRVGQ ELLPDMPRCVNPSRFNTARISDVYWAFYILLFGLTTSISIGIVERGWKNRLAIKRKILKNIL HRK >TbIr17 MHLYILIFIMQFLSLFSLEDTLFVINYFDIRKIRNVVGFTCNQDAQKTLTDIKMFSYFNKTI RISKLSADQISYLFLLKHSQHRLGIFVDKECMEDDLLIPFFNESTKLYLYTFKYKWLIISTD LDTTLQKLDDYHFGIMTDLIIAVPQSDANQYFLYEVFNPCRYRGAELVVNQIGDWTDNQ FNASFFDQDFPRRYDLGGIVLKMSGLTVYRPPDTLVSKYLQASQNISKDSMTKFTYTLTL HYADFFNFSLDVGDYTKWSPEDRVGPVVHGLDDRSIDICGSPLIMTEERAKYVEFLFPIW PFRTFFLFRTTLTSHIRLSEFLLPFMKLCWYWLIAITFISSFVLSFILTFEHTKLSTVDIVGA GLVITLGSICQQGSAFILQKTPGRIAFLSELILSLLIYNYYSASVVSTRLDKPIDKINDSLVE LAKTDIKMAAEPSLFFNFVLNKSEWEVNIFRKYWYMIPENQRFLVPEKAVKQIHKGGFA YHTTPEAIYPLVERTFGNRKICELTEVHLSRPQFLYINTRLDSPFTEIIRLGSVIMHERGLL KRQLIRWQTRRPECRTDVLSAESITMIETAPSFFLLIFGMFLSLIILIIENVVYRKSKIHKMK NQKKKRNPIIRRAKTNWLMKINKIKTI >TbIr18 MTKYSYAVALHWADMFNFTVDVHDYIEWKAGDLVGPVVHGLDSRDIDISGSPTIMTHG RAKYVDYIYHIWPYKSFFLFRTPSSSHIRMGEFLAPFAINCWYIVVFLCVVATVIFGVVLA FENMHISKIELYTIGLIITTGSLCQQGSAYIPAKAPGRIAFLNTMIFSLLLYNYYSASVVSSR LDKPIDKMNDSLVELSKTDLALAAEPTLFFDFQLNISDWEISIFRKKWYKTPINARFIFPEE AAKKIYKGGFAYHTTPEATYPYIERTFDNRRICELTEVRMGRPGYLYLNTRINSPFTEIAR LGSIKMNENGLIRRQLIRWKTRKPECRKDILSAASITMIEVAPCFCLWILGIILSLIIFELEK LVFMISHGKQIKLRRNKNSNLALKASWLKTTFQLK >TbIr19 MNTLNLCIKMIKYTLLFFACTILQIPILHCLEDCSFLVSYFEAKRVSSIVGFTCNKTDQTKF VETKFFSRYKNFIKLLNLEPESIAVLHFLEKANRNLGIFVDRKCADNSMFNQFYSESNRL FLFTQMYKWLIISTDLDRSVKRLNDSTFSITTDLTIAVFHKNDYVLHEAYNPCKYRKGRL FINEIGKWKAGVFVNVTFFDKNFIRRFNLEGMVVKTAGYNDYRPKDVDVLEYMLDTRD

457 KTKDSFTKFAIVLLLQLTEIFNFTLDIKDYTTWGDTGMLSPAVVGLFNQSIDISGNAVLMI EPRMVYVNYVFPMWPFRTFFIFRTTSTSHVNMGEFLRPFHKSCWYLTGLLLITAMSVFV VTLTFEDISIRRVDILGQGFLLAVGSLSQQGSAYIPTKSPGRIAFLYIMLFSLLIFNYYSASV VSSRFSKPIDKINDSLVELLKTKVKLGAEKFAFFGPLINVKCSLLSNFIDYLCLTDKTDLFL FRDVSQQSEWETTQFRKVWESIPDSEKILPKEEGLRRVERGGFAYHTRPESAYPTIERTF DNRKICELTEVQLRGSPYLYLATAKESPFEEITRLGSIKLAESGVWKRLLSKWTPRKLKC RRDILTSASIKMNEVVPCFVLLIAGVFLAVIIFLLELLVYSDHRGLRMSAKLESLKLVWW RYRMRS >TbIr20 LDKSVRRIDDSSFSITTDLIIAVFKTNEYVLHEAYNPCKYRDGRLFVNELGKWKGGGFD DIKFFDNNLPKRLNMGGMVLKTAGLNTYRPKDVDVREYMLDDRDKTKDSFTKFTCILL LHFTDLFNFTLAITDYSSWGDTGSLGPTVVNLANRTIDFNGNTVIMTKDRARCANFLFPI WPLRTFFILRTTLSSHIRMGEFLMPFDQSCWFSTALLLVIAIGAFIVTLNFEFIILTKIHVFG EGFLMAVGSLCQQGSAYIPRRSPGRIVLIQFMIFSLLIFNYYSASVVSSRINKPIDKINDSL VELLKTKVKVAAEPFAYFDFFLNQDGDWEISQFREHWLTKPENEKIMMLEEGIKRVKK GGFAYHTQQESAYPIIEKTFDNRKICELTEVHLVRPTGMFLVTAKNSPFEEMARYGSIKL AENGVWKRLLSKWVPRKLQCRKDILSSASIKMNEAIPQFALLLSGIFLSVLVLLIEKLVH GRHKGRRMSTNLEVLKLMWCNRNNKKVIE >TbIr21 MKHLLCSVCFVFICYIKIIRSNVPMQIFQSFVNLDQLNIVQVFHCWPHNEVLSSMKALRY KRHQLHNIQTGNIDLGQILHVNYYKVGILFDLDCERQISVTEQMQQNNLIFNETYHWLM YTTDQNCTNFLDGLPLSIDAEVTLAICDENCTLHDVYNPSFKHGGSLHIDLIGHWTQGGG LKMIDRSYKYDRRKNFDGIQLNFSTVIHNPPYGLDIISYLTKPIDPHIDTMQRYNFNLLLE LRDQYNFSINLIRYSTWGYLINGKFNGIIGDMINGVIDVAAVPMNIHKAERHAVMDYTV VIWTVKLSIFFRHPKKNLMKNNFLKPLSESVWYAALFVTAANWILLYATTKIRINYLKIR DDDKRESPFSDVSLITAAAVSQQGAYEVSGILSYKIIFLSLFTWAMLVYQFYSASIVGSLI AEPPRFITNVYELADSPLEFGAEDIGYMIDLFDTSTDSALRYLYDKKVSAHKKKPKKLFY SVEEGFKKVKKGGFAFYLDTATAYKTIQETFSDDEICELSELEIIKPVQCSTATNKFSPFN EMITYGLRKMVERGSQKRLKNVWYHKRPQCPESHKSKPTAITLQEFFPAIIIVIGGVIGSA VIGIIEVIIHNYKRRSVVA >TbIr22 MKRLVFSYFSLVCGMSYELDVNFLKDFYNNKNINQIVWFGCYTNSEVVSFVKDLSDSG FRVGYFDITDDNVKMKETLKVNYYKIGVILNDDCPDSQMIFDKFTGDKLVYNESYNWLI FTTDMIETKQIITKLPMNIATECTIAYKIGNNISLFEVYNPSYKHKANVNITHMGEWTKKS GLNYLYSDYKYVRRANLHGLKLNLSVFANNLPFGVPLDDYLRIPKKRHLDTMIRNNYA LLLQLKDLYNFTINLQKSTTWGYIINGTFNGVIGDMIKGIVDVSVAPMMIRPERLDVCEF TVQSYLMRGSIFLRHPKKNSIRNVFLTPFTEKAWFAVIGAGLIYWTVLLIIVIIEKKHTDN RMNVLHKRPFTNCIFTTVAALTQQGVTDVPRLYAGRIMCISLFIWSLLLFQFYSASIVGSL LAEQPKFINSIKDLVNSNIQVGIHDIAYMHNLFKMNTDPIIKELERKKINPNEKRKRSNYL NVTEGLSRVKTGGFAFYSDTATAYRIIQTTFTESEICDLVEIQLHPARDIGTVLPKKSPFK KMFIYGLRKIIEGGMTKRLYRIWHHQRPECPESFSSTPLPVHLQEFSPAVFLLAMGIGMAI IMLVAEIFYQKTIYFKRFNLDDLKILSN >TbIr23 MNRLIFSHFFLLIKFIIGYEMEVNFLKDYYEKKSINQIVWFACFTNPDMVSVVKVLSTSKF RVSYFDVTTTRTKIKESLKINYYKIGVVLNHDCSKSERIFDRFTADGLVYNDSYNWLIFT SNMNDLRKSISELPINVATEFTIANEMGNHISLFEVYNPSYKHNAITNITYMGKWDKKNG

458 LSFVSSEYKYVRRANLQGISLNFSVITYNDPSNIQLEEYLRVPKKRHLDTMSRKNYNMLL QLRDLYNFSITLKKATSWGYMVNGTFNGILGDMIKGIVDGSVAPLMIRPERLSVCEYTV QTYLMRGSIFLRHPKKNNIRNVFLTPFQDKAWLAVLGVAIIYCIVLLIIVRVEKNHTNKM SLLYKRPFTESAFTTIIALTQQGVTDVPWFYAGHIMCLSLFLWGLLLFQFYSASIVGSLIA EQPRFINTLRDLVESNLDVGIHDIAYMHNMFKTNPDPDVKDLGRIKIMPSAKRTKANFF NTTDGLNRVRAGGFAFYSDTSIAYRIIKTTFSEREICELVEIELHSPKHLSTVLPKNSPFKK MFIYGLRKIVESGMSKRLYRLWVHQRPKCPESYSSTPLPVQLQESSPALFLLIMGISIAIVI LLSEIFYQRKMKSVK

459