Insecta: Hymenoptera)

Systematics and evolution of the superfamily Platygastroidea (Insecta: Hymenoptera) 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 wasps. Most species of Platygastroidea are egg parasitoids of insects and spiders, or egg-larval parasitoids of gallflies, and many are recognized as important biological control agents of insect 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 wasp Trissolcus basalis. Calliscelio is a cosmopolitan genus of parasitoids that attack the eggs of crickets (Orthoptera: 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 taxonomy 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 Trissolcus basalis. 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 bees and sphecid wasps (Hymenoptera: Apoidea). 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, Platygastridae, Scelioninae). ZooKeys 730: 85–120. Zhu C, Luo S. Zhou X, Chen H, and Zhang G (2017) Earth BioGenome Project and Taxonomy. Biodiversity 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, Stephanidae). ZooKeys 612: 113–123. Chen H, Hong C, van Achterberg C, and Xu Z (2016) Description

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