Citation
Building a Diagnostic Framework for the Genus Synopeas Foerster (Hymenoptera: Platygastridae: Platygastrinae) Based on Reared Specimens from Papua New Guinea

Material Information

Title:
Building a Diagnostic Framework for the Genus Synopeas Foerster (Hymenoptera: Platygastridae: Platygastrinae) Based on Reared Specimens from Papua New Guinea
Creator:
Awad, Jessica Nemr
Publisher:
University of Florida
Publication Date:
Language:
English

Thesis/Dissertation Information

Degree:
Master's ( M.S.)
Degree Grantor:
University of Florida
Degree Disciplines:
Entomology and Nematology
Committee Chair:
Hodges,Amanda C
Committee Co-Chair:
Cave,Ronald David
Committee Members:
Talamas,Elijah Jacob
Graduation Date:
8/8/2020

Subjects

Subjects / Keywords:
coi
hymenoptera
microphotography
morphology
papua-new-guinea
parasitoid
platygastridae
platygastroidea
systematics
taxonomy

Notes

General Note:
The genus Synopeas Foerster (Hymenoptera: Platygastridae: Platygastrinae) has been largely neglected by modern professional systematics, and the absence of a diagnostic framework constitutes a serious taxonomic impediment. This study applied an integrative taxonomic approach to the problem by combining morphological, molecular, and ecological data to examine the bases for species identification in Synopeas. A non-destructive DNA extraction method allowed for the simultaneous analysis of COI barcode sequences and morphology for specimens reared from plant galls in Papua New Guinea. Important morphological characters included cuticular sculpture, setal arrangement, and ocular-ocellar length. The shape of the mesoscutellar spine was useful in some cases. The material yielded 17 new species of Synopeas, two previously described species, and one species that will not be described from a single damaged specimen. Most species were associated with one genus or species of gall host plant. Phylogenetic analysis supported the utility of COI for Synopeas species identification. The tools and procedures applied here constitute a diagnostic framework suitable for future revisionary work on Synopeas in other geographic regions.

Record Information

Source Institution:
UFRGP
Rights Management:
All applicable rights reserved by the source institution and holding location.
Embargo Date:
8/31/2022

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BUILDING A DIAGNOSTIC FRAMEWORK FOR THE GENUS SYNOPEAS F Ö RSTER (HYMENOPTERA: PLATYGASTRIDAE: PLATYGASTRINAE) BASED ON REARED SPECIMENS FROM PAPUA NEW GUINEA By JESSICA N . AWAD A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2020

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© 2020 Jessica N. Awad

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To my Grandma , Donna Suter Delong

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4 ACKNOWLEDGMENTS I would like to thank my supervisory committee chair, Dr. Amanda Hodges, for her professional advice and unwavering support throughout my degree program. I also thank my other committee members : Dr. Ronald D. Cave for his keen editing eye and scientific insight ; and Dr. Elijah Talamas for his guidance and mentorship in all things Platygastroidea. I feel very lucky to have had the opportunity to study with these three outstanding entomologists . I am grateful to my collaborators: Dr. Philip T. Butterill (Czech Academy of Sciences) for generating a wonderful collection of Synopeas , and for allowing me to conduct research on it ; an d Mr. Matthew R. Moore (Florida Department of Agricul ture) for providing DNA sequencing services and advice on analysis. This research was made possible by specimen loans from Dr. Lars Vilhelmsen (Natural History Museum of Denmark), Dr. Frederique Bakker (Naturalis Biodiversity Center Leiden), and Dr. Zoltán Vas (Hungarian Museum of Natural History). Mr. Charles Etienne Ferland (University of Guelph) and Mr. Derek Binns (UK) also generously contri buted Synopeas specimens to th e project. I thank the Florida Department of Agriculture, Division of Plant Industry, and the Florida State Collection of Arthropods for institutional support throughout my graduate studies . For their technical assistance at D PI , I am grateful to Ms. Johanna Schwartz, Mr. Jonathan Bremer, Mr. Charles Whitehill, Mr. Kyle Schnepp, Ms. Minjin Hao , and Ms. Cheryl Roberts . I than k librarian Mr. Jeff Eby for his help with locating and digitizing literature , and botanist Dr. Patti J. Anderson for her friendship and for sharing her expertise on tropical plants.

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5 For advice and encouragement, I thank Dr. Norm Leppla, Dr. Akito Kawahara , Dr. Emily Ellis, Dr. Jackie Miller, Dr. Debbie Matthews, the International Society of Hymenopterists, the Society of Systematic Biologists, the Center for Systematic Entomology, the McGuire Center for Lepidoptera and Biodiversity, the U niversity of Florida Linnaean team, the Biosecurity Research and Extension La b oratory , and my fellow entomology graduate students, who se company has greatly enriched my educational experience . I am much indebted to the technical and administrative staff of the Entomology and Nematology Department, especially Ms. Jenny Carr, Ms. Rut h Brumbaugh, Ms. Elena Alyanaya, Mr. Angel Perez, and Ms. Nancy Sanders, without who se kind assistance this work would not have been possible. I am particularly grateful to Dr. Y. Miles Zhang, who mentored me during his postdoc at the University of Florida and patiently fielded many questions about molecular analysis, phylogenetic software, gall biology, and general microhymenopteran matters . The Smithsonian Institut HYM Course sparked my interest in parasitoid taxonomy, for which I thank instructors Dr. Mike Gates, Dr. John Lill, Dr. Matt Buffington, and Dr. Bob Kula , as well as Ms. M ariel Guala and Ms. Amy Howe, who encouraged and enabled me to attend . I also thank Team Tephritid: Dr. Gary Steck, Dr. Marc Branham, Mr. Lou Somma, Mr. Erick Rodriguez, and Ms. Jessica Diaz, for providing employment during the last year of my studies and allowing me to hone my morphology skills on micrographs of fruit fly larvae. Finally, I would like to t hank my partner, Jim Hayden, for his support and encouragement throughout my graduate schooling, especially while I was finishing my thesis in quarantine during the COVID 19 pandemic .

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6 TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................ ................................ ................................ .. 4 LIST OF TABLES ................................ ................................ ................................ ............ 8 LIST OF FIGURES ................................ ................................ ................................ .......... 9 LIST OF ABBREVIATIONS ................................ ................................ ........................... 12 ABSTRACT ................................ ................................ ................................ ................... 13 CHAPTER 1 INTRODUCTION AND LITERATURE REVIEW ................................ ..................... 14 Gall Midges of Economic Importance ................................ ................................ ..... 14 Parasitoids of Gall Midges ................................ ................................ ...................... 16 Economic Importance of Synopeas ................................ ................................ ........ 17 Taxonomic Status of Synopeas ................................ ................................ .............. 22 Taxonomic Impediments to the Study of Synopeas ................................ ................ 25 COI Barcoding and Parasitoid Taxonomy ................................ ............................... 27 Objectives and Hypotheses ................................ ................................ .................... 28 2 SPECIES DESCRIPTIONS ................................ ................................ .................... 30 Introduction ................................ ................................ ................................ ............. 30 Materials and Methods ................................ ................................ ............................ 31 Material Examined ................................ ................................ ............................ 31 Morphological Techniques ................................ ................................ ................ 32 Analysis ................................ ................................ ................................ ............ 36 Species Descriptions ................................ ................................ .............................. 38 Synopeas sp. 1, new species ................................ ................................ ........... 38 Synopeas sp. 1b, new species ................................ ................................ ......... 40 Synopeas sp. 2, new species ................................ ................................ ........... 42 Synopeas sp. 3, new species ................................ ................................ ........... 45 Synopeas sp. 4, new species ................................ ................................ ........... 47 Synope as sp. 5, new species ................................ ................................ ........... 49 Synopeas sp. 6, new species ................................ ................................ ........... 51 Synopeas sp. 7, new species ................................ ................................ ........... 54 Synopeas sp. 8, new species ................................ ................................ ........... 56 Synopeas sp. 9, new species ................................ ................................ ........... 58 Synopeas sp. 9b, new species ................................ ................................ ......... 61 Synopeas sp. 10, new species ................................ ................................ ......... 63 Synopeas sp. 11, incertae sedis ................................ ................................ ....... 65 Sy nopeas sp. 13, new species ................................ ................................ ......... 67

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7 Synopeas sp. 15 (cf. Synopeas psychotriae Buhl) ................................ ........... 69 Synopeas sp. 16 (cf. Synopeas butterilli Buhl) ................................ ................. 72 Synopeas sp. 19, new species ................................ ................................ ......... 74 Syno peas sp. 22, new species ................................ ................................ ......... 76 Synopeas sp. 23, new species ................................ ................................ ......... 78 Synopeas sp. 24, new species ................................ ................................ ......... 80 Synopeas pterocarpi Buhl ................................ ................................ ................ 82 Synopeas csoszi Buhl ................................ ................................ ...................... 84 Key to the Species of Synopeas of New Guinea ................................ .................... 86 Discussion ................................ ................................ ................................ .............. 90 3 PHYLOGENETIC ANALYSIS ................................ ................................ ............... 101 Introduction ................................ ................................ ................................ ........... 10 1 Materials and Methods ................................ ................................ .......................... 102 Material Examined ................................ ................................ .......................... 102 Molecular Techniques ................................ ................................ .................... 102 Analysis ................................ ................................ ................................ .......... 104 Results ................................ ................................ ................................ .................. 105 Discussion ................................ ................................ ................................ ............ 111 4 CONCLUSIONS ................................ ................................ ................................ ... 114 Study Significance ................................ ................................ ................................ 114 Future Directi ons ................................ ................................ ................................ .. 115 LIST OF REFERENCES ................................ ................................ ............................. 117 BIOGRAPHICAL SKETCH ................................ ................................ .......................... 125

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8 LIST OF TABLES Table page 1 1 Species of Synopeas reared from agricultural pests. ................................ ......... 21 1 2 Number of species descriptions in the genus Synopeas by biogeographic region and year. Compiled from Hymenoptera Online (2019) . ........................... 24 2 1 Summary of Synopeas specimens from the Butterill loan to the Florida State Collection of Arthropods, by gall host plant iden tity. ................................ ........... 32 2 2 Morphological characters on which morphotypes of Synopeas are based . ........ 33 2 3 Species of Synopeas described from Australasia, Indonesia, and the Philippines. ................................ ................................ ................................ ......... 37 2 4 Known geographic distributions of gall host plants for Synopeas specimens from the Butterill loan to the Florida State Collection of Arthropods. .................. 94 3 1 Gall host plant associations of Synopeas mor phospecies sampled for COI barcode analysis. ................................ ................................ .............................. 107

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9 LIST OF FIGURES Figure page 1 1 Specimen of Synopeas subgenus Sactogaster , photographed after non destructive DNA extraction. Collected in Valles Caldera National Preserve, Sandoval County, New Mexico, September 2008, by staff of the Systematic Entomolog y Laboratory, United States Department of Agriculture. Photo by Jessica Awad. ................................ ................................ ................................ ..... 25 2 1 Two characters of the mesosoma used to separate morphotypes of Synopeas . A) Mesoscutellar spine. B) Pronotal microsculpture. Photos by Jessica Awad. ................................ ................................ ................................ ..... 36 2 2 Three characters of the frons used to separate morphotypes of Synopeas. A) Distance between lateral ocellus and compound eye. B) Central keel. C) Head macrosculpture. Photos by Jessica Awad. ................................ ................ 36 2 3 Synopeas species 1. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ............ 40 2 4 Synopeas species 1b. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 42 2 5 Synopeas species 2. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Ph otos by Jessica Awad. ............ 44 2 6 Synopeas species 3. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ............ 46 2 7 Synopeas species 4. Clockwise from top left: lateral habitus, ventral habitus, dor sal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ............ 49 2 8 Synopeas species 5. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ............ 51 2 9 Synopeas species 6. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ............ 54 2 10 Synopeas species 7. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ............ 56 2 11 Synopeas species 8. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ............ 58 2 12 Synopeas species 9. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ............ 61

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10 2 13 Synopeas species 9b. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 63 2 14 Synopeas species 10. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 65 2 15 Synopeas species 11. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 67 2 16 Synopeas species 13. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 69 2 17 Synopeas species 15. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 71 2 18 Synopeas species 16. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 74 2 19 Synopeas species 19. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 76 2 20 Synopeas species 22. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 78 2 21 Synopeas species 23. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 80 2 22 Synopeas species 24. Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ ................................ ................................ ................. 82 2 23 Holotype of Synopeas pterocarpi . Clockwise from top left: lateral habitus, anterior head, posterodorsal habitus, dorsal habitus. Scale bar represents 0.1 mm. Photos by Elijah Talamas. ................................ ................................ .... 84 2 24 Holotype of Synopeas csoszi . Scale bar represents 0.5 mm. Photo by Elijah Talamas. ................................ ................................ ................................ ............. 86

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11 2 25 Comparative morphology of the second metasomal sternite (S2). Clockwise from top left: Synopeas spp. 1, 1b, 2, 3, 6, and 13. Photos by Jessica Awad. ... 95 2 26 Comparative morphology of the second metasomal tergite (T2). Clockwise from top left: Synopeas spp. 1, 1b, 13, and 9. Photos by Jessica Awad. ........... 96 2 27 Comparative morphology of the forewing. Clockwise from top left: Short and sparse setae with fringe uniform in length ( Synopeas sp. 16); Long and sparse setae with fringe longer on posterior margin ( Synopeas sp. 24); Short and dense setae, proximally glabrous with linea setosa ( Synopeas sp. 2); Short and dense setae with fringe longer on posterior margin ( Synopeas sp. 13). Photos by Jessica Awad. ................................ ................................ ............. 97 2 28 Inostemma specimen reared from cecidomyiid gall on Macaranga . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. ................................ .................... 98 2 29 Holotype of Synopeas quasimodo . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Photos by Elijah T alamas. ................................ . 99 2 30 Comparative morphology of the mesoscutellar spine. Top row: absent or inconspicuous. Center row: short and pointed. B ottom row: long and blunt. Photos by Jessica Awad. ................................ ................................ .................. 100 3 1 Neighbor joining tree of COI barcode data from Synopeas of New Guinea, inc luding Platygaster and Inostemma species as outgroups, and Synopeas specimens from the UK, the USA, and Myanmar. ................................ ............ 108 3 2 Maximum likelihood analysis of COI barcode data from Synopeas of New Guinea, including Platygaster and Inostemma as outgroups, and Synopeas specimens from t he UK, the USA, and Myanmar. ................................ ............ 109 3 3 Maximum parsimony bootstrap consensus tree of COI barcode data from Synopeas of New Guinea, incl uding Platygaster and Inostemma as outgroups, and Synopeas specimens from the UK, the USA, and Myanmar. .. 110

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12 LIST OF ABBREVIATIONS BIN Barcode Index Number BMNH British Museum of Natural History, London BOLD Barcode of Life Database COI Cytochrome oxidase I CUID Collection unit identi fi er DPI Division of Plant Industry FDACS Florida Department of Agriculture and Consumer Services FSCA Florida State Collection of Arthropods GTR+F+R3 General time reversible model with empirical base frequencies and three rate categories HNHM Hungarian Museum of Natural History, Budapest HOL Hymenoptera Online MUSCLE Multiple Sequence Comparison by Log Expectation NA Not applicable NBCL Naturalis Biodiversity Center, Leiden OOL Ocular ocellar length PCR Polymerase chain reaction PNG Papua New Guinea POWO Plants of the World Online Database SAM South Australian Museum s.s. Sensu stricto TBR Tree Bisection Regrafting USNM United States National Museum, Washington ZMUC Zoological Museum, University of Copenhagen

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13 Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science BUILDING A DIAGNOSTIC FRAMEWORK FOR THE GENUS SYNOPEAS F Ö RSTER (HYMENOPTERA: PLATYGASTRIDAE: PLATYGASTRINAE) BASED ON REARED SPECIMENS FROM PAPUA NEW GUINEA By Jessica N. Awad August 2020 Chair: Amanda Hodges Major: Entomology and Nematology The genus Synopeas F ö rster (Hymenoptera: Platygastridae: Platygastrinae) has been largely neglected by modern professional systematics, and the absence of a diagnostic framework constitutes a serious taxonomic impediment. Th is study applie d an integrative taxonomic approach to t he problem by combining morphological, molecular, and ecological data to examine the bases for species identification in Synopeas . A non destructive DNA extraction method allowed for the simultaneous analysis of COI b arcode sequences and morphology f or specimens reared from plant galls in Papua New Guinea . Important morphological characters included cuticular sculpture, setal arrangement, and ocular ocellar length. The shape of the meso scutellar spine was useful in som e cases . The material yielded 17 new species of Synopeas , two previously described species, and one species that will not be described from a single damaged specimen . Most species were associated with one genus or species of gall host plant. Phylogenetic a nalysis supported the utility of COI for Synopeas species identification. The tools and procedures applied here constitute a diagnostic framework suitable for future revisionary work on Synopeas in other geographic regions .

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14 CHAPTER 1 INTRODUCTION AND LITERATURE REVIEW Gall Midges of Economic Importance Cecidomyiidae, generally referred to as gall midges, comprise 6 , 590 described species in 812 genera (Gagn é and Jaschhof , 2017). Approx imately one fourth of all known cecidomyiid species are fungivores in the early diverging subfamilies Catotrichinae, Lestremiinae, Micromyinae, Winnertziinae, and Porricondylinae. These fungivores have little to no economic significance, with the exception s of Camptomyia corticalis (Loew) and C. heterotobia Mamaev , which are pests of shiitake mushrooms in East Asia (Shin et al . , to members of the subfamily Cecidomyiinae, which includes the plant gall makers and other herbivores. Cecidomyiinae are the largest subfamily of Cecidomyiidae, having diversified with flowering plants during the Cretaceous Period. Th is subfamily include s nearly all plant pest species for the family , and the subfamily also includes fungivorous and predatory species. Some cecidomyiine predators are considered beneficial natural enemies of minute agricultural pests such as mites, aphids, and scale insects (Gagn é and Jaschhof , 2017). Other herbivorous gall midges are used as biological control agents of invasive weeds (Dorchin , 2008). C ecidomyiid pest species that attack grain have the potential to cause serious economic losses in developed areas, and severe food shortages fo r subsistence farmers in developing nations. The Hessian fly, Mayetiola destructor (Say) , is a well known pest of wheat in western Asia, North Africa, Europe, and North America (Shukle , 2009). Other pests of wheat include the saddle gall midge , Haplodiplos is marginata (von

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15 Roser) , the wheat midge , Sitodiplosis mosellana (Gehin) , and the lemon wheat midge , Contarinia tritici (Kirby) . The African rice gall midge, Orseolia oryzivora Harris and Gagné , can cause 25 to 50% yield loss by preventing panicle formati on (Nwilene et al . , 2006). In Asia, rice is similarly threatened by the rice gall midge , O. oryzae Wood Mason (Samalo et al . , 1983). Fruit crops are also susceptible to damage from gall midges. Several species of Dasineura Rondani specialize on different genera of fruit trees and bushes. Dasineura mali (Kieffer) is a pest of apple in New Zealand, Europe, and North America. Although it attacks the leaves rather than the fruit, D. mali can impact the long term yield potential of a tree, and cause quarantine issues for importing regions (He and Wang , 2011). Larvae of the blueberry gall midge, D. oxycoccana (Johnson) , feed within flower buds, causing up to 80% loss of flower production (Sampson et al . , 2006). D asineura amaramanjarae Grover attack s the flowers o f mango. The leaves and flowers of mango also host many other species of gall midge in the genera Procontarinia Kieffer and Cecconi, Gephyraulus Rübsaamen, and Asynapta Loew . Although most of the mango midge species originate in southern Asia, members of this pest complex sometimes become a problem in other mango growing regions (Pena et al ., 1998). Most recently, P. mangiferae (Felt) was detected in Cuba, and it may spread t o other parts of the Caribbean and southern Florida (Rodriguez Tapia et al . , 2018). Besides causing direct damage to marketable plant parts and indirect damage via reduction in photosynthetic ability, some species of Cecidomyiidae have been implicated in i ncreased susceptibility to plant pathogens. Infestation of mango foliage by Procontarinia mangicola (Shi) leads to secondary damage by the anthracnose fungus

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16 Colletotrichum gloeosporioides (Pena et al . , 1998). In Cyprus, Dasineura oleae (Angelini) is consi dered a minor pest of olive, but it appears to promote susceptibility to the olive knot bacterium Pseudomonas savastanoi (Iordanou , 1982). It is certain that other associations between gall midges and plant pathogens remain to be discovered in agricultural systems. Parasitoids of G all M idges Specialist parasitoids have evolved their own strategies for overcoming the challenge of the cecidomyiid plant gall. Some attack eggs or young larvae while others oviposit through the gall to reach older larvae. Most g all midge parasitoids are found in the h ymenoptera n superfamilies Chalcidoidea, Ceraphronoidea, and Platygastroidea. Within Chalcidoidea, most belong to Eulophidae (Nwilene et al . , 2006 ; Sampson et al . , 2002 ; Pena et al . , 1998). Notable examples include Aprostocetus procerae (Risbec) on O . oryzivora , Tetrastichus Haliday species on D . oxycoccana and P. mangiferae , and Closterocerus pulcherrimus (Kerrich) on Procontarinia matteiana Kieffer and Cecconi . Closterocerus pulcherrimus is native to south ern Asia, but adventive in South African mango orchards, where it provides some control (Morgan et al . , 2017). Other important chalcidoids are in the eupelmid gen us Eupelmus Dalman , and the pteromalid genera Systasis Walker and Macroglenes Westwood . All three genera are found in Indian mango systems, while Macroglenes penetrans (Kirby) is an egg parasitoid of S . mosellana in Canadian wheat (Pena et al . , 1998 ; Smith et al . , 2004). Within Ceraphronoidea, species of Aphanogmus Thoms on and Ceraphron pallidiventris Ashmead are abundant in cranberry and blueberry crops, but the impact of their activity is unknown (Sampson et al . , 2002).

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17 Within Platygastrida e , the subfamily Platygastrinae is comprised of parasitoids of gall midges. The genus Platygaster Latreille contains the largest number of species of importance to gall midge control, although this genus is not a natural group (Elijah Talamas, per s. comm. ). Platygaster demades Walker is native to Europ e and was introduced to New Zealand, where it causes significant reductions in populations of D . mali (He and Wang , 2011). Platygaster species are also implicated in control of P . tumifex in balsam fir, O . oryzivora in African rice, O . oryzae in Asian rice , M . destructor in wheat, D . oxycoccana in blueberry, and Dasineura species in mango (Kelley , 2009 ; Nwilene et al , 2006 ; Samalo et al . , 1983 ; Schmid et al . , 2018 ; Sampson et al . , 2006 ; Pena et al . , 1998). The platygastrine genera Synopeas F ö rster and Inostemma Haliday also parasitize gall midges of economic importance. Synopeas temporale Austin is the most abundant parasitoid of Procontarinia matteiana in Pakistan . U nidentified Synopeas and Inostemma species parasitized a significant percentage of gall midges in Mississippi blueberries (Mahmood Ur Rehman et al . , 2013 ; Sampson et al . , 2006). Economic Importance of Synopeas Platygastroidea are the third largest superfamily of parasitoid wasps, exceeded in number of described species only by Ichneumonoidea and Chalcidoidea , and include s approximately 6 , 0 00 described species. Historically , the superfamily consist ed of two families, Scelionidae and Platygastridae (Austin et al ., 2005). They are small wasps (usually less than 3 mm long) with reduced wing venation. The toruli are located low on the f rons , less than their own diameter from the clypeus (Mason , 1993). Pla tygastroidea are characterized by the specialized structure of the metasoma and ovipositor. The ovipositor is retracted inside the metasoma, often with a telescoping mechanism formed

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18 of intersegmental conjunctiva that allow s for maximum extension of the ov ipositor. I nternal coiling mechanisms of the ovipositor and external projections or extensions of the metasoma, such as horns, humps, or sac like structures , are common . The metasomal tergites and sternites are typically appressed to allow for ovipositor e xtension via hydrostatic pressure, and the metasomal spiracles are reduced or absent in most lineages . The superfamily is also characterized by the presence of papillary ( basiconic ) sensilla, also called multiporous gustatory sensilla, on the distal antenn omeres in the female (Masner , 1993). The ground plan biology of Platygastroidea is considered to be endoparasitism of insect eggs (Austin et al ., 2005). Platygastridae comprise approximately 2 , 1 00 species in about 7 0 genera. There are two subfamilies: Scel iotrachelinae and Platygastrinae. Sceliotrachelinae include egg insects such as Coccoidea and Aleyroidea (Austin et al. , 2005) . As far as is known, Platygastrinae consist entirely of gall midge parasitoids (Masner , 1993 ; Austin et al ., 2005). The biology of Platygastrinae has not been well explored relative to their diversity , in part due to their small size and concealment within hosts that are themselves concealed feeder s. Female platygastrines oviposit in eggs or larvae of the host. In some species, w asp eggs or first instar s persist in a state of delayed development until the host reaches the prepupal stage, at which point the parasitoid development resumes and proceeds rapidly (Kim et al ., 2011 ; Ogah et al ., 2010). Many species are solitary, but others are gregarious, and some are polyembryon ic (Austin et al ., 2005 ; Clausen , 1978 ; Sampson et al ., 2006). In solitary species, physical combat may remove supernumerary larvae (Abram et al ., 2012a).

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19 Platygaster is the largest genus in Platygastrinae and has the most species associated with gall midges of economic importance (Hymenoptera Online , 2019). H owever, molecular evidence does not support the monophyly of Platygaster , so these numbers do not reflect meaningful biological reality (Elijah Talamas, pers. comm.). Indeed, Platygaster is traditionally diagnosed by a lack of distinguishing characters (Wa lker , 1873). Synopeas is monophyletic according to a recent phylogeny based on ultraconserved elements ( Bonnie Blaimer, pers . comm. ). The genus can be diagnosed morphologically, most obviously by dense setation on the posterior mesosoma and anterior metas oma that obscur es the junction between metasomal tergites 1 and 2 (Figure 1 1) . Metasomal tergites 1 and 2 are fused, sternites 1 and 2 are fused, and the forewings do not have a long fringe of setae on the posterior margin. Synopeas has a cohesive generic concept, while Platygaster is a poorly defined genus, and the number of Platygaster species make it unwieldy for revisionary work at the student level. After Platygaster , the genus Synopeas contains the most species of interest to the biological control o f Cecidomyiidae. In the southeastern United States, the blueberry gall midge , D. oxycoccana , and the blueberry tip midge, Prodiplosis vaccinnii (Felt) , are attacked by two unidentified species of Synopeas . These species were recogniz ed as the most common parasitoids in Mississippi blueberries, with higher parasitism rates than by Platygaster , Inostemma , and Aprostocetus Westwood (Hymenoptera: Eulophidae) in the same system. Sampson et al . (2006) discovered that Synopeas females enter blueberry buds and oviposit near the midgut of the first instar midge. They illustrated the immature stages throughout

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20 development and sequenced COI barcodes, but no species names were attached due to a lack of identification resources. Synopeas myles (Wal ker) is well studie d due to its association with C ontarinia nasturtii in Europe. It arrived adventitiously in North America in the first decade of the 2000s and was first detected in Canada in 2015 (Mason et al ., 2017). It is the most abundant parasitoid o f C. nasturtii in Switzerland , but it was never considered for deliberate introduction due to its apparently wide host range. However, these host records were based on literature, so it i s possible that the references did not correct ly identif y the parasit oid. Of the six reports of S. myles attacking species other than C. nasturtii , only one was vouchered. Due to the taxonomic impediments to Synopeas identification, unvouchered reports may not be considered reliable. New experiments are needed to confirm th e host range of S. myles (Abram et al ., 2012b). In southern Asia, three Synopeas species are associated with the mango midge pest complex. Synopeas procon Austin and S. temporale were first reared from P . matteiana , and S. mangiferae Austin was reared from Procontarinia echinogalliperda (Mani) (Austin , 1984). Mahmood et al . (2013) reported that S. temporale was the most abundant parasitoid of P. matteiana in Pakistan and that the phenology of host and parasitoid were tightly synchronized. Levels of host specificity within this system remain uncertain. There has been little exploration into the natural enemies of D . amaramanjarae and other Dasineura species that form galls in reproductive structures of mango (Mahmood Ur Rehman et al ., 2014). Many S ynopeas species have been reared from agricultural pests but have not undergone further pest management research (Table 1 1 ) . Other species of Synopeas

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21 have been recovered from agricultural crops, but with no information on the insect host. For example, S . otiosum Kieffer (1924) was reared from banana in Algeria. Synopeas osgoodi MacGown (1974) was reared from fir trees in Maine, where the balsam gall midge is a pest of Christmas tree farms. In general, it is much easier to record host plant information tha n host insect information. T he plant is much larger than the insect; the plant remains identifiable after galling and parasitism; and the taxonomy of plants, particularly those of economic importance, is better resolved than that of the Cecidomyiidae. Of t he 378 valid species of Synopeas , host information is available for only 42 species. Among the 336 species with no ecological data, there are likely to be other parasitoids of potential economic importance. One area that merits special attention is the ide ntity and prevalence of Synopeas species attacking beneficial gall midges. The only record found of such was S . oleae Buhl and Viggiani reared from Lasioptera berlesiana Paoli, a natural enemy of the olive fly , Dacus oleae Gmel in (Diptera: Tephritidae) (Bu hl and Viggiani , 2008). Table 1 1 . Species of Synopeas reared from agricultural pests . Synopeas species Insect host Plant host Reference S. craterum (Walker) Resseliella ribis (Marikovskij) Blackcurrant Walker, 1836 S. rufipes (Ashmead) Contarinia gossypii Felt Cotton Ashmead, 1893 S. curvicauda (F ö rster) Contarinia pisi (Loew) Pea Foerster, 1856 S. lugubre Thomson Dasineura napi (Loew) Cole crops Thomson, 1859 S. daucicola Kieffer Kiefferia pericarpiicola (Bremi) Carrot Kieffer, 1916 S. talhouki Vlug Odinadiplosis amygdali (Anagnostopoulos) Almond Vlug, 1976 S. procon Austin S. temporale Austin Procontarinia matteiana Kieffer and Cecconi Mango Austin, 1984 S. mangiferae Austin Procontarinia echinogalliperda (Mani) Mango Austin, 1984 S. laurae (Vlug) Contarinia tritici (Kirby) Wheat Vlug, 1991 S. myles (Walker) Contarinia nasturtii (Kieffer) Cole crops Abram et al ., 2012b

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22 Taxonomic S tatus of Synopeas The genus Synopeas was propos ed by F ö rster in 1856. The original diagnosis is Die Unterrandader an der Spitze ohne Knöpfchen; Das Schildchen mehr oder weniger verlängert, nicht halbkreisig, oder wenn verkurzt, dann immer von der Seite zusammangedrückt, mit einer pfriemen oder warzenför migen Spitze versehen; Der Hinterleib nicht besonders verlängert; Das zweite Bauchsegment beim nicht sackartig [ Submarginal vein without pterostigma; scutellum more or less extended, not semicircular, or if shortened, then always laterally compressed , with pointed or wart shaped process; abdomen not especially extended; second abdominal segment of female not sac like . ] Interes tingly, there is no mention of the setation on the anterior metasoma and petiole that is characteristic of all known species of Synopeas . After the description of Synopeas , some species of Platygaster and Epimeces Westwood were transferred to Synopeas . Th us, the earliest species descriptions predate F ö Synopeas ventrale (Westwood) in 1833. Nees von Esenbeck described three species in 1834 that would later be placed in Synopeas . N o hymenopteran taxonomy would be complete without a generous contribution from Francis Walker, who described 12 species in 1836 (one only as a figure) and another five species in 1839. Masner (1964) list ed S. inermis Thomson as the type species, after Muesebeck (1951). Vlug ( 199 5) lis t ed Platygaster sosis Walker as the type species, but his reasoning is unclear, as no type species was designated by F ö rster (1856). Ashmead (1893) designate d S. prospectus F ö rster as the type species. must take prior ity.

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23 In his 1856 work, F ö rster described several other platygastrid genera, including Sactogaster , which was distinguished from Synopeas Das zweite Bauchsegment beim stark zusammengedrückt, sackartig [ segment of the female compressed, sac ] . Kozlov (1978) treated Sactogaster as a subgenus of Synopeas , and this opinion was seconded by Vlug (1995). The ventral expansion of sternite 2 t hat accommodate s the ovipositor appears to be the only character distinguishing Sactogaster from the rest of Synopeas ( Figure 1 ) . The subgenus contains 21 species. The genus Dolichotrypes was propos ed by Crawford and Bradley in 1911. Like Sactogaster , its distinguishing character is the shape of the female metasoma. In Dolichotrypes , the metasoma is extremely elongate, primarily through the elongation of metasomal segments 4 through 6 . It was synonymized with Synopeas by Masner (1964). F our species were described in Dolichotrypes from 1911 to 1966. Fouts (1924) synonymiz ed Synopeas with Leptacis F ö rster, based on the presence of four clavomeres in the female antenna. This opinion was s har ed by Muesebeck (1951) and has caused a good deal of confusion. In F ö the original work in which both genera were described, Leptacis is distinguished from Synopeas based on characters of the scutellar spine (F ö rster , 1856). Masner (1960 ; 1964) restored Synopeas as a valid genus , describing, for the first time, the transverse the same piles at the base of the first metasomal segment. However, some confusion remains, particularly when authors are working with outdated references. As recently as

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24 200 7, nine new species of Leptacis from India were described, all of which were subsequently transferred to Synopeas (Ushakumari and Narendran, 2007; Buhl, 2009). Of the 378 valid species of Synopeas , 124 described species are from the Palearctic , more than a ny other biogeographical region (Table 1 2 ). Prior to 1990, the Palearctic and Nearctic regions yielded the most species descriptions. After 1990, the rate of species descriptions increased, with the Palearctic (68) and Indomalayan (65) regions leading in number, followed by the Afrotropic al (46) and Neotropic al (42) regions. The Australasian and Oceanic regions have the fewest described species of Synopeas , although the number of Australasian descriptions increased after 1990. The Nearctic is the only regi on in which the number of new Synopeas species descriptions decreased after 1990. Table 1 2 . Number of species descriptions in the genus Synopeas by biogeographic region and year. Compiled from Hymenoptera Online (2019). Ecozone Number of species described before 1990 Number of species described after 1990 Total Afrotropic al 12 46 58 Australasia 4 18 22 Indomalaya 9 65 74 Nearctic 38 6 44 Neotropic al 12 42 54 Oceania 1 1 2 Palearctic 56 68 124 Total 132 246 378

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25 Figure 1 1 . Specimen of Synopeas subgenus Sactogaster , photographed after non destructive DNA extraction. Collected in Valles Caldera National Preserve, Sandoval County, New Mexico, September 2008, by staff of the Systematic Entomology Laboratory, United States De partment of Agriculture. Photo by Jessica Awad. Taxonomic I mpediments to the S tudy of Synopeas Since the 1990s, the taxonomic impediment to Synopeas identification has become increasingly severe . Of the 246 species described after 1990, the majority are the work of a single prolific hobbyist who is not formally trained . The descriptions are scattered throughout many publications that are difficult to access by conventional means. There is no phyl ogenetic context, no molecular data, no ecological data, and the images provided are insufficient for diagnosis. The illustrations are generally simple line drawings, often excluding the mouthparts, frontal view of the head, and lateral and

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26 dorsal views of the mesosoma. These l ine drawings do not depict microsculpture, which is an important source of characters. The probability of unreliable characters occurring in prior work is high. Many species descriptions after 1990 rely heavily on measurements taken from one or a few specimens . Parasit ic wasps vary in size depending upon host identity, host age, nutritional quality , and other developmental factors (Godfray , 1994 ; Quicke, 1997 ). Morphometric measurements do not necessarily scale isometrically with size . In fact, there is much evidence for allometric scaling in parasitoid morphology (Janzon, 1986; Parreno et al. , 2017 ; Souza et al. , 2018; Goltz et al ., 2020 ) . For example, the platygastroid Telenomus alsophilae Vier e ck exhibits differences in body size and antennal proportions when reared from different sizes and species of host egg (Johnson et al. , 1987). Ther efore, the measurements of one individual cannot be assumed to represent the measurements of the entire species . Morphometrics can generate reliable characters if taken from many individuals sampled from a range of ecological conditions. In this way, varia tion within a species may be distinguished from variation between species . Ultimately, the holotypes of each of these putative species must be imaged and catalogued, as the descriptions are minimally useful for identification. To begin resolving the taxono mic issues with Synopeas , it is necessary to gather morphological, molecular, and ecological data. The importance of characters for species diagnosis can be more appropriately ascertained once relevant data have been obtained and synthesized.

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27 COI B arcoding and P arasitoid T axonomy COI barcoding was proposed by Hebert et al. (2003) as a global identification system for animals. The system was designed as a taxonomic aid and identification tool, especially for morphologically constrained or otherwise challenging taxa and life stages. The mitochondrial gene cytochrome c oxidase I (COI) exhibits a high rate of molecular evolution, few indels, no introns, and a haploid mode of inheritance , providing a strong phylogenetic signal. In addition, the primers f or COI are reliable , widely available, and generally considered to be the barcode standard of choice for zoology (Taylor and Harris, 2012) . Using lepidopteran examples, Hebert et al. ( 2003 ) demonstrated that COI barcoding could be useful for species delimi tation, discovery of cryptic species, and clarifying the taxonomy of poorly known groups. Since 2003, DNA barcoding has spawned hundreds of research papers and more than a million records in online databases , with a strong taxonomic bias towards arthropods (Taylor and Harris , 2012). As barcoding technology has achieved widespread usage, its limitations have also become apparent. Analytical issues may affect the interpretation of genetic distance between sampling units (Rubinoff et al . , 2006). Endosymbionts and parasitic microbes can influence selection on mitochondria, leading to unexpected diversity or lack of diversity in COI (Hurst and Jiggins , 2005). Barcoding is not suitable for species delimitation in taxa with high intraspecific variability in the barcode region , as is the case with some Diptera (Meier et al ., 2006). P arasit ic Hymenoptera , on the other hand, show high congruence between COI sequence data and morphologically recognizable taxa (Quicke et al. , 2012).

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28 Despite its limi tations, COI barcoding is a valuable part of an integrative taxonomic approach, in which molecular data is not the sole source of diagnostic characters but is complementary to morphological, ecological, physiological, and biogeographic data (Gokhman , 2018) . Thus, DNA barcoding and other PCR based techniques remain relevant to the development of parasitoid diagnostics. Potential applications include elucidat ing host parasitoid interactions, delimitation of cryptic species and strains, and enhance d monitoring and evaluation of introduced biological control agents (Gariepy et al. , 2007). Such advanced applications first require the establishment of a robust diagnostic framework. Objectives and Hypotheses Th e current project applies an integrative taxonomic approach to build a diagnostic framework of useful characters for the identification of Synopeas species. Th e overarching goal can be divided into two primary objectives: Primary objective 1 to identify morpholog ical and ecological characters that are associated with genetic differences ; Primary objective 2 to identify morphological and ecological characters that vary within a species, i.e., characters that are not useful for diagnosis. The secondary goal of thi s project is to use the data to elucidate species relationships within the examined material. This goal can be divided into four secondary objectives: Secondary objective 1 to construct a phylogeny of operational taxonomic units ; Secondary objective 2 to delimit species based on totality of evidence ; Secondary objective 3 to attach appropriate taxonomic n ames to the examined material ; Secondary objective 4 to create a morphological key to the examined species.

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29 I hypothesize that some morphological v ariation occurs within a single species, and that some characters will prove unreliable for diagnosis.

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30 CHAPTER 2 SPECIES DESCRIPTIONS Introduction The described Synopeas species of Papua New Guinea can be traced to three collecting events: the voyage of the Noona Dan in 1961 1962 (Wolff, 1963; Buhl, 1997); a visit to Mt. Wilhelm by the Hungarian acarologist János Balogh in 1968 (Dózsa Farkas, 2003; Buhl, 2004); and the extensive r earing work of Philip Butterill in 2010 2012. Of these, only Balogh and Butterill sampled the island of New Guinea. The Noona Dan expedition was sponsored by the University of Copenhagen. Its crew included zoologists, botanists, geographers, and anthropol ogists. Its mission was to collect natural history specimens and data from several Indo Australian island groups. The Papuan Synopeas specimens described from this voyage were collected from the Bismarck Archipelago, off the island of New Britain. The Noon a Dan expedition also collected Synopeas specimens from the Philippines and the Solomon Islands, which I examined and compared to the material from the Butterill collection. The Noona Dan specimens are deposited in the Zoological Museum of the University o f Copenhagen. The present work describes the Synopeas species from the island of New Guinea. The se include 17 new species of Synopeas from the Butterill collection and four previously described species from the Butterill and Balogh collections : S. butterilli Buhl, S. csoszi Buhl, S. psychotriae Buhl, and S. pterocarpi Buhl . A key to the Synopeas species of New Guinea is provided.

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31 Materials and Methods Material Examined The material used for this study is likely rich collection of Synopeas specimens, in terms of ecological and molecular information potential. The parasitoid collection was loaned to Elijah Talamas (Florida State Collection of Arthropo ds) by Philip T. Butterill, who studied the ecology of gall forming insects in Papua New Guinea from 2010 to 2012 (Butterill and Novotny, 2015). All specimens were accompanied by detailed ecological data and stored in 95% ethanol, making them suitable for molecular analysis. In total, the collection includes 520 platygastrine specimens from 146 galls, reared from 20 plant genera in 16 families. Of these, 245 specimens are identifiable as Synopeas (Table 2 1) , reared from 17 plant species in 12 genera . All S ynopeas emerged from cecidomyiid galls, except for the gall on Aglaia rimosa (Meliaceae ), which was associated with Psylloidea (Hemiptera). P syllids and cecidomyiids can be found in the same gall (Mani, 1964) , and so I do not consider this to be evidence that these Synopeas are psylloid parasitoids . Inostemma and Platygaster specimens from the Butterill collection were selected as outgroups . For geographic reference, the analysis also include d Synopeas specimens from Myanmar, the United Kingdom, and the United States. Holotypes of new Synopeas species are to be deposited in the United States National Museum of Natural History in Washington, DC. Paratypes will be deposited in the Florida State Collection of Arthropods in Gainesville, Florida.

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32 Table 2 1 . Summary of Synopeas specimens from the Butterill loan to the Florida State Collection of Arthropods, by gall host plant identity. Host plant family Host plant species Number of galls Number of specimens Actinidaceae Saurauia conferta 5 25 Saurauia poolei 1 1 Saurauia schumanniana 1 2 Elaeocarpaceae Elaeocarpus sp. 1 1 1 Euphorbiaceae Homalanthus nervosus 1 1 Macaranga strigosa 3 9 Loganiaceae Neuburgia corynocarpa 2 13 Meliaceae Aglaia rimosa 1 1 Myristicaceae Paramyristica cf. sepicana 1 3 Piperaceae Piper amboinense 7 14 Piper celtidiforme 2 5 Rubiaceae Nauclea sp. 1 41 111 Nauclea tenuiflora 8 24 Psychotria ramuensis 1 8 Urticaceae Cypholophus sp. 1 4 22 Cypholophus friesianus 2 5 Total 12 genera 81 245 Morphological Techniques Initial hypotheses of species concepts were based on external morphology examined with light microscopy (Table 2 2, Fig ure 2 1, Fig ure 2 2). Images that clearly illustrate diagnostic characters were made of each morphotype by using microphotographic techniques. Microphotography hardware consisted o f a Macropod® imaging system with a Canon EOS 6D Mark II camera, EF 70 200mm lens, and 20X M Plan APO Mitutoyo objective lens. Microphotography software included EOS 6D Mark II camera utility, Helicon Focus Pro 6.8.0 for image stacking, and Adobe Photoshop for addition of scale bars and post processing. The d orsal and lateral habitus and frontal view of the head of each morphotype were imaged, as well as other characters of interest.

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33 The color of the body, legs, and mesoscutellar spine w ere included as mor phological character s because they ha ve been previously used to describe Synopeas species. However, cuticular color is not recommended for species delimitation, as it is often variable and may be affected by specimen age, condition, and preservation technique . Table 2 2 . Morphological characters on which morphotypes of Synopeas are based . Character group Character Character states Head Shape in anterior view 1. Circular 2. Distinctly ovoid 3. Triangular 4. Slightly ovoid Central keel 1. Complete 2. Absent 3. Partial 4. Impressed Sculpture on frons 1. Rugose macrosculpture 2. Reticulate microsculpture 3 . Rugulose macrosculpture Epi torular sculpture 1. Rugulose 2. Reticulate microsculpture 3. Rugose Number of clypeal setae 1. Two 2. Four L engths of median pair of clypeal setae 1. Longer than lateral pair 2. Equal to lateral pair Arrangement of clypeal setae 1. Evenly spaced 2. Medial setae closer to each other than to lateral setae 3. Medial setae closer to lateral setae than to each other Shape of mandible 1. Bidentate 2. Unidentate Distance between lateral ocellus and compound eye (OOL) 1. Less than one ocellar diameter 2. Approximately one ocellar diameter 3. Greater than one ocellar diameter Hyperoccipital carina 1. Complete 2. In terrupted medially 3. Present only in medial third 4 . Absent Distance between lateral ocellus and hyperoccipital carina 1. Less than one ocellar diameter 2. Approximately one ocellar diameter 3. Greater than one ocellar diameter

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34 Table 2 2. Continued. Mesosoma Epomial carina 1. Present , extending to dorsal limit of pronotum 2. Absent 3. Present only in ventral half Microsculpture of lateral pronotum 1. Present throughout 2. Present dorsally, absent ventrally 3. Present medially, absent dorsally and ventrally 4. Present anterodorsally, absent posteroventrally 5. Entirely or almost entirely absent 6. Present dorsally, absent ventrally, with smooth band anterior to tegula Mesoscutellar spine 1. Absent or inconspicuous 2. Sh ort and pointed 3. Long and blunt 4. Broadly triangular 5. Long and sharp S cutellar spine in lateral view 1. Pointing posteriorly 2. Angled posterodorsally Margin of p ropodeal keel in lateral view 1. Rounded 2. Pointed ventrally 3. Pointed dorsally and ventrally 4. Posteriorly flattened Mesosomal dorsum in lateral view 1. Flat 2. Weakly convex 3. Convex 4. Strongly convex Notauli 1. Distinctly grooved 2 . Unmarked or faintly suggested Parapsidal lines 1. Very faint 2. Unmarked 3. Marked by absence of sculpture 4. Well impressed Setation of mesoscutum 1. Very dense 2. Sparse 3. Moderately dense 4. A bsent medially , present laterally Mesos cuta l lamella 1. Long and narrow 2. Short, projecting less than its own width beyond margin 3. Broad and rounded 4. Short and narrow 5. Roughly triangular 6. Broad and square 7. Truncate, with striate sculpture 8. Defined by carinae

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35 Table 2 2. Continued. Setation of mesoscutellum 1. Present throughout 2. Absent medially 3. Present lateral to striate sculpture 4. Laterally dense, medially sparse Metasoma Microsculpture of S 2 1. Narrow band at posterior margin 2. Present at posterior margin and in posterolateral corners 3. Present on posterior half 4. Absent 5. Present in posterolateral corners Sculpture of T2 1. Absent 2 . Wide transverse band of microsculpture at posterior margin 3. Narrow transverse band of microsculpture at posterior margin 4. Longitudinally s triate 5. Microsculpture present in patch on posterior margin, not reaching posterolateral corners Length of T2 1. Shorter than mesosoma 2. Approximately as long as mesosoma 3. Longer than head and mesosoma combined Wing Length of setae on disc of forewing 1. L onger than distance between setal bases 2. S horter than distance between setal bases Density of setae on disc of forewing 1. Dense 2. Sparse Arrangement of setae on disc of forewing 1. Uniformly setose distally, proximally glabrous with l inea setosa 2. Setose distally, glabrous proximally 3. Entirely setose Forewing setal fringe 1. Setae on posterior margin distinctly longer than setae on anterior margin 2. Approximately uniform in length on anterior and posterior margins Color Body color 1. Black 2. Dark brown 3. B rown Color of legs 1. Coxae brown, otherwise yellow 2. Coxae brown, otherwise yellow to brown Color of scutellar spine 1. Concolorous with mesoscutellar disc 2. Yellow

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36 Figure 2 1 . Two characters of the mesosoma used to separate morphotypes of Synopeas . A) Mesos cutellar spine. B) Pronotal microsculpture. Photos by Jessica Awad. Figure 2 2 . Three characters of the frons used to separate morphotypes of Synopeas. A) Distance between later al ocellus and compound eye. B) Centr al keel. C) Head macrosculpture. Photos by Jessica Awad. Analysis A character matrix of morphological data was constructed in the online vSysLab (Johnson, 2008) . A morphological identification key was created. Names w ere assigned to specimens by comparison to holotypes of Australasian and S outheast Asian species (Table 2 3). If no matching types were found , species were described as new.

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37 Table 2 3 . Species of Synopeas described from Australasia , Indonesia, and the Philippines . Species Year Type locality Holotype location Host plant genus S. isus (Walker) 1839 Tasmania BMNH Unknown S. leda (Walker) 1839 Tasmania BMNH Unknown S. idarniforme (Dodd) 1916 Australia SAM Unknown S. saccharale (Dodd) 1916 Australia SAM Saccharum S. decumbens Buhl 1997 Papua New Guinea ZMUC Unknown S. lemkaminensis Buhl 1997 Papua New Guinea ZMUC Unknown S. saintexuperyi Buhl 1997 Papua New Guinea ZMUC Unknown S. ventricosum Buhl 1997 Papua New Guinea ZMUC Unknown S. acutiventris Buhl 1997 Philippines ZMUC Unknown S. balabacensis Buhl 1997 Philippines ZMUC Unknown S. crassiceps Buhl 1997 Philippines ZMUC Unknown S. luteolipes Buhl 1997 Philippines ZMUC Unknown S. montanus Buhl 1997 Philippines ZMUC Unknown S. mukerjeei Buhl 1997 Philippines ZMUC Unknown S. palawanensis Buhl 1997 Philippines ZMUC Unknown S. pallescens Buhl 1997 Philippines ZMUC Unknown S. csoszi Buhl 2004 Papua New Guinea HNHM Unknown S. pleuralis Buhl 2004 Australia HNHM Unknown S. queenslandicus Buhl 2004 Australia HNHM Unknown S. achterbergi Buhl 2008 Indonesia NBCL Unknown S. ciliaris Buhl 2008 Indonesia NBCL Unknown S. flavispinum Buhl 2008 Indonesia NBCL Unknown S. halmaherense Buhl 2008 Indonesia NBCL Unknown S. infuscatus Buhl 2008 Indonesia NBCL Unknown S. lombokensis Buhl 2008 Indonesia NBCL Unknown S. longulum Buhl 2008 Indonesia NBCL Unknown S. nigricoxa Buhl 2008 Indonesia NBCL Unknown S. praemorsus Buhl 2008 Indonesia NBCL Unknown S. quasimodo Buhl 2008 Indonesia NBCL Unknown S. novaezealandiae Buhl 2011 New Zealand ZMUC Unknown S. butterilli Buhl 2013 Papua New Guinea USNM Paramyristica S. psychotriae Buhl 2013 Papua New Guinea USNM Psychotria S. pterocarpi Buhl 2013 Papua New Guinea USNM Pterocarpus S. alternatum Buhl 2014 Australia ZMUC Unknown S. bangmadseni Buhl 2014 Australia ZMUC Unknown S. carlseni Buhl 2014 Australia ZMUC Unknown S. eucalypti Buhl 2014 Australia ZMUC Unknown S. salicorniae Buhl 2014 Australia ZMUC Unknown S. striolagaster Buhl 2014 Australia ZMUC Unknown S. triangulatum Buhl 2014 Australia ZMUC Unknown BMNH: British Museum of Natural History, London. SAM: South Australian Museum, Adelaide. ZMUC: Natural History Museum of Denmark, Copenhagen. HNHM: Hungarian National History Museum, Budapest. NBCL: Naturalis Biodiversity Center, Leiden. USNM: United State s National Museum, Washington, DC.

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38 Species Descriptions The present study describes 17 new Synopeas species and redescribes four species from the holotypes , applying the unified species concept ( de Quieroz , 2007). The unified concept defines a species as a (monophyly, diagnosability, reproductive isolation) are considered as different properties tha t accumulate during lineage separation. Each property is important but not solely necessary to the species concept. Thus, the present work integrates multiple lines of evidence (molecular, morphological, and ecological) to support the designation of specie s . Special emphasis was placed on morphological diagnos is. Synopeas sp. 1 , new species Description . Body length 1.1 1.4 mm. B ody black , coxae brown , otherwise legs yellow to brown . Head s lightly ovoid. Central keel a bsent , micros culpture on frons r eticulate , e pi torular microsculpture reticulate , sometimes with minute rugulae. Four clypeal setae , medial setae longest , m andible bidentate . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina robust , complete. Distance between lat eral ocellus and hyperoccipital carina less than one ocellar diameter . Mesosoma . Epomial carina present , complete or nearly so . L ateral pronot al microsculpture present dorsally, smooth ventrally. Mesoscutellar spine short and pointed , pointing posteriorly in lateral view . P ropodeal keel pointed ventrally in lateral

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39 view . Mesosomal dorsally convex in lateral view . Notaul i unmarked or faintly suggested. Parapsidal lines very faint. Seta e sparse o n mesoscutum . Mesoscutal lamella oblong. Seta e o n me soscutellum laterally dense, medially sparse. Metasoma . Microsculpture on S 2 present at posterior margin and in posterolateral corners . Sculpture on T2 present in wide transverse band at posterior margin . T2 shorter than or equal to length of mesosoma. Forew ing. Setae on disc short , dense . D isc uniformly setose distally , proximally glabrous with linea setosa . S etal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis . Synopeas sp. 1 has a short, pointed scutellar spine and a robust , complete hyperoccipital carina. It may be distinguished from similar species by the patch of microsculpture on posterior and posterolateral S2 (Figure 2 25 ) and the wide band of reticulate microsculpture on dorsal T 2 (Figure 2 26 ) . It resembles Synopeas sp. 1b but differs in the length of T 2, which is much more elongate in Synopeas sp. 1b ( c ompare Figures 2 3 and 2 4 ) . Synopeas sp. 1 also has sparser setation on the frons than does Synopeas sp. 2 ( c ompare Figures 2 3 and 2 5 ) . Plant Associations . Reared from blister galls on Saurauia conferta and Saurauia schumanniana (Actinidaceae). Material E xamined . 7 females, 5 males. Papua New Guinea, Morobe Province, Yawan , 6 .1414142 °S , 146.8741517 °E , Coll. P. Butterill . Ex. cecidomyiid galls. 2 females and 2 males emerged 17 V 2012 from Saurauia conferta . 1 female emerged 03 IX 2012 from Saurauia schumanniana . 1 female emerged 03 IX 2012 from Saurauia conferta . 3 females and 3 males emerged 05 XI 2012 from Saurauia conferta .

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40 Figure 2 3 . Synopeas sp ecies 1 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 1 b , new species Descrip tion. Body length 1.1 1.5 mm. Body black , coxae brown , legs otherwise yellow to brown . Head in frontal view slightly ovoid. Centra l keel absent , micro s culpture on frons reticulate , e pi torular sculpture reticulate. Clypeal setae unknown, mandible unknown . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina robust , complete. Distance between lateral ocellus and hyperoccipital carina less than one ocellar diameter .

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41 Mesosoma. Epomial carina present, complete or nearly so . L ate ral pronot al microsculpture present dorsally, smooth ventrally. Mesoscutellar spine short and pointed , pointing posteriorly in lateral view . Propodeal keel pointed ventrally in lateral view. Mesosoma dors ally convex in lateral view. Notauli unmarked or fai ntly suggested. Parapsidal lines very faint. Seta e sparse o n mesoscutum. M esoscutal lamella short, projecting less than its width beyond margin. Seta e o n mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S2 present at posterior margin and in posterolateral corners . Microsculpture on T2 present in narrow band at posterior margin . T2 approximately as long as mesosoma . Forew ing. Setae on disc short , dense . D isc uniformly setose distally, proximally glabrous with linea seto sa. S etal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. Synopeas sp. 1 b has a short, pointed scutellar spine and a robust , complete hyperoccipital carina. It may be distinguished from similar looking species by the patch of microsculpture on posterior and posterolateral S2 (Figure 2 25 ) . It resembles Synopeas sp. 1, but differs in the length of T 2, which is much more elongate in Synopeas sp. 1b ( c ompare Figures 2 3 and 2 4 ) . Additionally, Synopeas sp. 1 has a wide patch of microsculpture at the dorsal margin of T2, which in Synopeas sp. 1b is much narrower (Figure 2 26) . Plant Associations. Reared from nodule like galls on Cypholoph u s friesianus (Urticaceae).

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42 Material E xamined . 6 females, 2 males. P apua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid galls on Cypholophus friesianus . 4 females emerged 04 X 2010. 2 males emerged 07 X 2010. 2 females emerged 24 V 2012. Figure 2 4 . Synopeas sp ecies 1b . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm . Photos by Jessica Awad. Synopeas sp. 2 , new specie s Description. Body length 1.2 1.5 mm. Body black , coxae brown, legs otherwise yellow to brown . Head in frontal view circular . Frontal keel absent , micro culpture on frons reticulate , e pi torular sculpture reticulate with minute rugulae. Four clypeal setae , medial

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43 setae longest , closer to each other than to lateral setae , mandible bidentate . Ocular oce llar length less than one ocellar diameter . Hyperoccipital carina robust , complete. Distance between lateral ocellus and hyperoccipital carina approximately one ocellar diameter . Mesosoma. Epomial carina present, complete or nearly so . Lateral pronotal mi crosculpture present dorsally, smooth ventrally. Mesoscutellar spine short and pointed , pointing posteriorly in lateral view . Propodeal keel rounded in lateral view. Mesosoma dorsally convex in lateral view . Notauli unmarked or faintly suggested. Parapsidal lines very faint. Seta e on mesoscutum medially absent, laterally present. Mesoscutal lamella broad and rounded or roughly triangular . Setae on mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S2 present in narrow band at posterior margin. M icrosculpture on T2 present in narrow band at posterior margin. T2 shorter than or equal to length of mesosoma. Forew ing. Setae on disc short , dense . D isc uniformly setose distally, proximally glabrous with linea setosa. S etal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. Synopeas sp. 2 has a short, pointed meso scutellar spine and a robust , complete hyperoccipital carina. The metasomal sculpture on T2 is restricted to a narrow band at the posterior ma rgi n (Figure 2 5 ) . It is similar to Synopeas sp. 24, and these can be differentiated by the hyperoccipital carina. In Synopeas sp. 24, the hyperoccipital carina is medially weakened and laterally present as a thin carina, whereas in Synopeas sp. 2, the hyperoccipital carina is uniformly robust.

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44 Plant Associations. Reared from blister galls on Piper amboinense and Piper celtidiforme (Piperaceae). Material E xamined . 6 females, 5 males. Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146. 8741517 °E , Coll. P. Butterill. Ex. cecidomyiid galls. 1 female and 1 male emerged 20 IX 2010 from Piper amboinense . 1 male emerged 0 5 X 2010 from Piper . 1 female emerged 12 X 2010 from Piper celtidiforme . 1 male emerged 18 X 2010 from Piper amboinense . 3 females emerged 15 X 2010 from Piper amboinense. 1 male and 1 female emerged 25 X 2010 from Piper amboinense . 1 male emerged 11 XI 2010 from Piper amboinense . Figure 2 5 . Synopeas sp ecies 2 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad.

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45 Synopeas sp. 3 , new species Description. Body length 1.1 1.3 mm. Body black or dark brown , legs brown. Head in frontal view slightly ovoid. Central keel partial and impressed , reticulate microsculpture on frons , e pi torular sculpture reticulate with minute rugulae. Four clypeal setae , equal in length, evenly spaced , mandible bidentate . Ocular ocellar length approximately one oce llar diameter . Hyperoccipital carina weak, complete. Distance between lateral ocellus and hyperoccipital carina less than one ocellar diameter . Mesosoma. Epomial carina present , complete or nearly so . L ateral pronot al microsculpture present dorsally, smoo th ventrally, with smooth band anterior to tegula. Mesoscutellar spine short and pointed , angled posterodorsally in lateral view . Propodeal keel pointed dorsally and ventrally in lateral view . Mesosoma dors ally convex in lateral view. Notauli unmarked or f aintly suggested. Parapsidal lines very faint , marked by absence of sculpture. Seta e sparse o n mesoscutum. M esoscutal lamella broad and rounde d or roughly triangular. Seta e o n mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S 2 present in narrow band at posterior margin. T2 smooth. T2 shorter than mesosoma . Forew ing. Setae on disc medium to long , dense . D isc uniformly setose distally, proximally glabrous with few scattered setae . S etal fringe uniform in lengt h on posterior and anterior margins. Diagnosis. Synopeas sp. 3 has a short, pointed mesoscutellar spine that is angled upward, and a weak but complete hyperoccipital carina. It resembles Synopeas sp. 24 and may be distinguished by the presence of a partial central keel on the frons. The forewing of Synopeas sp. 3 has setae of approximately equal length around the

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46 wing margin, while in Synopeas sp. 24, the setae are distinctly longer on the posterior margin (Figure 2 27) . Plant Associations. Reared from circular galls on Neuburgia corynocarpa (Apocynaceae). Material E xamined . 1 female, 4 males. Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Emerged 22 X 2012 from cecidomyiid gall on Neuburgia corynoca rpa . Figure 2 6 . Synopeas sp ecies 3 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad.

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47 Synopeas sp. 4 , new species Description. Body length 1.1 1.5 mm. Body black , coxae brown, legs otherwise yellow to brown . Head in frontal view circular. Central keel complete , micros culpture on frons reticulate , e pi torular sculpture reticulate with minute rugulae. Four clypeal setae , medial longest , closer to each other than to lateral setae , m andible bidentate . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina weak, complete. Distance between lateral ocellus and hyperoccipital carina approximately one ocellar diameter . Mesosoma. Epomial carina present , complete or nearly so . Lateral pronotal m icrosculpture present dorsally, smooth ventrally. Mesoscutellar spine short and pointed , angled posterodorsally in lateral view . Propodeal keel pointed ventrally in lateral view. Mesosoma dors ally convex in lateral view . Notauli unmarked or faintly suggested. Parapsidal lines very faint. Seta e sparse o n mesoscutu m . M esoscutal lamella roughly triangular. Mesoscutellum medially glabrous , laterally setose . Metasoma. Microsculpture on S 2 present in narrow band at posteri or margin. Microsculpture on T2 present in very narrow band at posterior margin. T2 shorter than or equal to length of mesosoma. Forew ing. Setae on disc short , sparse , sparser proximally than distally . S etal fringe somewhat longer on posterior margin than on anterior margin. Diagnosis. Synopeas sp. 4 has a strong, distinct central keel on the frons. Synopeas sp. 15 also has a line on the frons that extends from between the toruli to the median ocellus, but it is impressed ra ther than raised. Additionally, the parapsidal lines of Synopeas sp. 15 are well impressed and are very weak in Synopeas sp. 4 (compare

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48 Figures 2 7 and 2 17) . Synopeas sp. 16 may have a complete central keel, but it is weakly developed. The sculpture of th e lateral pronotum in Synopeas sp. 16 is restricted to a narrow medial band (Figure 2 18) , rather than covering the dorsal half of the lateral pronotum as in Synopeas sp. 4. Plant Associations. Reared from node like and blister galls on Nauclea ( Rubiaceae). Occasionally reared from the same gall as Synopeas sp. 19. Material E xamined . 5 females, 6 males. Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid galls on Nauclea sp. 1. 1 male emerged 1 1 XI 2010. 1 male emerged 25 XI 2010. 1 male emerged 26 XI 2010 . 2 female s emerged 01 XII 2010 . 1 female emerged 26 I 2011. 2 females and 3 males emerged 21 XI 2012.

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49 Figure 2 7 . Synopeas sp ecies 4 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. S ynopeas sp. 5, new species Description. Body length 1.3 1.5 mm. Body black , coxae brown, legs otherwise yellow to brown. H ead in frontal view large and triangular. Frontal keel absent, but angle of frons gives impression of a keel in certain lights , micro culpture on frons reticulate , e pi torular sculpture reticulate. Four clypeal setae , medial longest , evenly spaced , mandible bidentate . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina absent.

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50 Mesosoma. Epomial carina present, complete or nearly so . Lateral pronotal microsculpture present medially, smooth ventrally , with narrow smooth area s dorsally and posteriorly . Mesoscutellar spine absent or inconspicuous. Propodeal keel rounded in lateral view . Mesosoma dors ally convex in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines very faint. Seta e moderately dense on mesoscutu m. M esoscutal lamella short, projecting less than its width beyond margin. Seta e on mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S 2 present in narrow band at posterior margin. T2 smooth . T2 shorter than or equal to length of mesosoma. Forew ing. Setae o n disc short , dense . D isc uniformly setose distally, proximally glabrous with linea setosa. S etal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. Synopeas sp. 5 has an exceptionally large, triangular head, unlike any other known species of Synopeas from the region . The malar space is long, about equal to the height of the compound eye, and the mesoscutellar spine is inconspicuous to absent . Comments. A similar head shape is found in Inostemma macarangae Buhl (Figure 2 28 ) , which has also been reared from Macaranga galls, suggesting convergent evolution. Plant Associations. Reared from round fuzzy galls on Macaranga strigosa (Euphorbiaceae).

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51 Material E xamined . 2 females. Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid galls. Emerged 21 XI 2012 from cecidomyiid gall on Macaranga strigosa . Figure 2 8 . Synopeas species 5 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 6 , new species Description. Body length 1.1 1.5 mm. Body black to dark brown , coxae brown , legs otherwise yellow to brown . Head in frontal view blocky . Central keel present ventrally , absent dorsally, micros culpture on frons reticulate , e pi torular microsculpture reticula te, sometimes with

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52 minute rugulae. Four clypeal setae , medial longest , closer to each other than to lateral setae ; mandible bidentate , somewhat short, tips just meeting in middle rather than overlapping . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina weak, medially present, laterally absent. Distance between lateral ocellus and hyperoccipital carina greater than one ocellar diameter . Mesosoma. Epomial carina absent or greatly reduced . L ateral pronot al microsculpture present thr oughout. Mesoscutellar spine inconspicuous. Propodeal keel rounded in lateral view. Mesosoma dors ally convex in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines unmarked. Seta e on mesoscutum medially absent, laterally present. M esoscut al lamella broad and rounded or roughly square. M esoscutellum medially glabrous , laterally setose . Metasoma. Microsculpture on S 2 present on posterior half. Microsculpture on T2 present in wide band at posterior margin. T2 shorter than mesosoma. Wing. Seta e of forewing disc short , dense . Forewing disc uniformly setose distally, proximally glabrous with linea setosa. Forewing setal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. Among the species treated here, Synopeas sp. 6 may be recognized by the absent or reduced epomial carina. When present, the epomial carina is short, extending less than halfway up the pronotum. The meso scutellar spine is absent or inconspicuous. S 2 is covered with reticulate microsculpture on the pos terior half (Figure 2 25) , and there is a wide band of microsculpture at the posterior margin of dorsal T2 (Figure 2 26) . The shape of the head in Synopeas sp. 6 is distinctly blocky in anterior view.

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53 Comments. The distance between the dorsal margin of th e eye and the vertex is very small, as in Synopeas sp. 5. It thus may represent an intermediate form between the typical head shape in Synopeas and the extreme found in Synopeas sp. 5. Plant Associations. Reared mainly from circular and blister galls on Na uclea (Rubiaceae) . One record from Homalanthus nervosus (Euphorbiaceae) . Material E xamined . 10 females, 4 males. Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid galls. 1 female emerged 24 VII 2010 from Nauclea sp. 1. 1 male emerged 16 XI 2010 from Nauclea sp. 1. 1 female emerged 17 XI 2010 from Homalanthus nervosus. 1 female emerged 25 XI 2010 from Nauclea sp. 1. 2 female s emerged 26 XI 2010 from Nauclea sp. 1. 5 female s, 3 males emerged 30 XI 2010 from Nauclea sp. 1.

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54 Figure 2 9 . Synopeas species 6 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 7 , new species Description. Body length 1.1 1.3 mm. Body black , coxae brown , legs otherwise yellow to brown . Head in frontal view distinctly ovoid. Central keel present ventrally, micros culpture on frons reticulate , e pi torular sculpture rugulose. Four clypeal setae , med ial longest , closer to lateral setae than to each other , mandible unidentate . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina absent. Mesosoma. Epomial carina present, complete or nearly so . Lateral pronotal microsculpture present dorsally, smooth ventrally. Mesoscutellar spine inconspicuous.

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55 Propodeal keel pointed dorsally and ventrally in lateral view. Mesosoma dors ally convex in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines very fain t. Seta e sparse o n mesoscutum. M esoscutal lamella roughly triangular. Seta e o n mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S 2 present in narrow band at posterior margin. T2 smooth, sometimes with narrow band of microsculptur e at posterior margin . T2 shorter than or equal to length of mesosoma. Forew ing. S eta e on disc moderately long , sparse . D isc uniformly setose distally, glabrous proximally. S etal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. Synopeas sp. 7 may be recognized by its head shape, which is distinctly wider than high, and by the absence of a hyperoccipital carina. It resembles Synopeas sp. 22, it but can be easily distinguished by the shape of the dorsal mesosoma, which is only moderately convex i n Synopeas sp. 7 and distinctly convex in Synopeas sp. 22 (compare Figures 2 10 and 2 20) . Additionally, the posterior vertex of Synopeas sp. 22 is medially depressed, while the vertex of Synopeas sp. 7 is not. Plant Associations. Reared from variously sha ped galls on Nauclea sp p. (Rubiaceae) and Macaranga strigosa (Euphorbiaceae). Material E xamined . 4 females, 11 males. Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid galls. 1 male emerged 15 XI 2010 from Nauclea sp. 1. 1 male emerged 16 XI 2010 from Nauclea sp. 1. 1 male emerged 24 I 2011 from Nauclea sp. 1. 2 females, 5 males emerged 25 I 2011 from Nauclea tenuiflora . 1 female, 1 male emerged 26 I 2011 from Macaranga

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56 strigo sa . 1 female, 1 male emerged 26 I 2011 from Nauclea sp. 1. 1 male emerged 08 X 2012 from Nauclea tenuiflora . Figure 2 10 . Synopeas species 7 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 8 , new species Description. Body length 0.9 1.25 mm. Body black , coxae brown, legs otherwise yellow to brown . Head in frontal view slightly ovoid. Central keel present ventrally , micros culpture on frons reticulate , e pi torular sculpture reticulate with minute rugulae. Four clypeal setae , medial longest , closer to each other than to lateral setae , mandibl e bidentate . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina medially

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57 strong, laterally weak or absent. Distance between lateral ocellus and hyperoccipital carina approximately one ocellar diameter . Mesosoma. Epomial carina pre sent , complete or nearly so . Lateral pronotal microsculpture present throughout. Mesoscutellar spine inconspicuous. Propodeal keel rounded in lateral view. Mesosoma dors ally convex in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines ve ry faint. Seta e moderately dense on mesoscutum. Mesoscutal lamella broad and square. Setae on m esoscutellum laterally dense, medially sparse . Metasoma. Microsculpture on S 2 present in narrow band at posterior margin. T2 smooth , with very narrow band of microsculpture at posterior margin . T2 shorter than or equal to length of mesosoma. Wing. Setae on forewing disc short , dense . Forewing disc uniformly smooth distally, proximally glabrous with linea setosa. Forewing setal fringe somewhat lon ger on posterior margin than on anterior margin. Diagnosis. Synopeas sp. 8 has the pronotum entirely covered with reticulate microsculpture , which is not typical among the species treated here . The absent or inconspicuous mesoscutellar spine also helps to identify it . It can be distinguished from the similar Synopeas sp. 6 by the epomial carina, which is complete or nearly so in Synopeas sp. 8 and abent or reduced in Synopeas sp. 6. Plant Associations. Reared from variously shaped galls on Nauclea sp. 1 (R ubiaceae). Material E xamined . 9 females, 1 male. Papua New Guinea, Morobe Province, Yawa n, 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid galls on

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58 Nauclea sp. 1 . 6 females emerged 15 XI 2010 . 1 female emerged 16 XI 2010. 1 female emerged 25 XI 2010. 1 male emerged 26 I 2011. 1 female emerged 04 X 2012. Figure 2 11 . Synopeas species 8 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 9 , new species Description. Body length 1.2 1.8 mm. Body black , coxae brown, l eg s otherwise dark yellow to brown. Head in frontal view distinctly ovoid. Central keel present ventrally, absent medially, sometimes also present do rsally, just ventral to median ocellus ; macros culpture on frons rugose , e pi torular sculpture rugose. Four clypeal setae , medial

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59 longest , closer to each other than to lateral setae , mandible bidentate . Ocular ocellar length approximately one ocellar diamete r . Hyperoccipital carina weak, incomplete, or obscured by rugose head sculpture. Distance between lateral ocellus and hyperoccipital carina approximately one ocellar diameter . Mesosoma. Epomial carina nearly complete , with posterior terminal pit . Lateral pronotal microsculpture present dorsally, smooth ventrally, with smooth band anterior to tegula. Mesoscutellar spine long and blunt, papilliform or wart like , pointing posteriorly in lateral view . Propodeal keel rounded in lateral view. Mesosoma do rs ally convex in lateral view. Notauli distinctly grooved . Parapsidal lines marked by absence of sculpture. Seta e sparse o n mesoscutum. Mesoscutal lamella truncate, with striate sculpture. M esoscutellum striate ly sculpture d , laterally setose . Metasoma. Mi crosculpture on S 2 present in narrow band at posterior margin. T2 smooth, with very narrow band of microsculpture at posterior margin . T2 shorter than or equal to length of mesosoma. Wing. Setae on forewing disc short , sparse , sparser proximally than distally . Forewing setal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. The head of this species is mostly covered in rugose macrosculpture and the notauli are distinctly grooved , characters which it shares with Synopeas spp. 9b and 19. The mesoscutellar spine of Synopeas sp. 9 is papilliform or wart like, with the length of the spine only slightly greater than its thickness in lateral view. Synopeas sp. 9b has a mesoscutellar spine that is similar in form, but its length is more than twice the thickness at its midpoint ( c ompare Figures 2 12 and 2 13) . The mesoscutellar spine in Synopeas sp. 19 is absent or inconspicuous (Figure 2 19) .

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60 Plant Associations. Reared from blister galls on Nauclea sp p . (Rubiaceae). Material E xamined . 12 females, 7 males. Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid galls. 1 female emerged 01 VII 2010 from Nauclea sp. 1. 1 female emerged 23 VII 2010 from Nauclea sp. 1. 1 female emerge d 24 VIII 2010 from Nauclea sp. 1. 2 females, 3 males emerged 22 IX 2010 from Nauclea tenuiflora . 1 fe male emerged 2 6 X 2010 from Nauclea sp. 1. 1 male emerged 30 X 2010 from Nauclea sp. 1. 1 male emerged 26 XI 2010 from Nauclea sp. 1. 1 female, 1 male emerged 29 XI 2010 from Nauclea sp. 1. 1 female emerged 26 I 2011 from Nauclea sp. 1. 1 female emerged 01 VIII 2012 from Nauclea sp. 1. 1 female emerged 21 XI 2012 from Nauclea sp. 1. 1 female emerged 17 X 2012 from Nauclea tenuiflora . 1 f emale, 1 male emerged 13 XI 2012 from Nauclea sp. 1. 1 female emerged 21 XI 2012 from Nauclea sp. 1.

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61 Figure 2 12 . Synopeas species 9 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 9b , new species Description. Body length 1.3 1.9 mm. Body black , coxae brown , legs otherwise dark yellow to brown . Head in frontal view ovoid. Centr al keel partial, present ventrally , absent medially, sometimes present dorsally ; macros culpture on frons rugose , e pi torular sculpture rug o se. Four clypeal setae , medial longest , closer to each other than to lateral setae , mandible bidentate . Ocular ocellar length greater than or equal to one ocellar diameter . Hyperoccipital carina absent .

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62 Mesosoma. Epomial carina complete or nearly so . Lateral pronot al microsculpture present dor sally, smooth ventrally. Mesoscutellar spine long and blunt , pointing posteriorly in lateral view . Propodeal keel pointed ventrally in lateral view. Mesosoma dors ally convex in lateral view. Notauli distinctly grooved . Parapsidal lines marked by absence of sculpture. Seta e sparse o n mesoscutum. M esoscutal lamella truncate, with striate sculpture. M esoscutellum striate ly sculpture d, laterally setose . Metasoma. Microsculpture on S 2 present in narrow band at posterior margin. T2 smooth, with medium to wide band of microsculpture at posterior margin . T2 shorter than mesosoma. Forew ing. Setae on disc short to medium in length, dense . D isc uniformly setose distally, glabrous proximally. S etal fringe distinctly longer on posterior margin than on anterior m argin. Diagnosis. The head of this species is mostly covered in rugose macrosculpture and the notauli are distinctly grooved , characters which it shares with Synopeas spp. 9 and 19. The mesoscutellar spine of Synopeas sp. 9 b is papilliform or wart like, wi th its length more tha n twice the thickness at its midpoint . Synopeas sp. 9 has a mesoscutellar spine that is similar in form, but the length is only slightly greater than its thickness in lateral view (compare Figures 2 12 and 2 13) . The mesoscutellar spine in Synopeas sp. 19 is absent or inconspicuous (Figure 2 19) . Plant Associations. Reared from blister gall on Nauclea sp. (Rubiaceae). Material E xamined . 1 female specimen . Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741 517 °E , Coll. P. Butterill. Ex. cecidomyiid galls on Nauclea sp. 1. Emerged 30 XI 2010.

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63 Figure 2 13 . Synopeas species 9b . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. S ynopeas sp. 10 , new species Description. Body length 1.6 mm. Body black , coxae brown, legs otherwise yellow to brown. Head in frontal view slightly ovoid. Centr al keel absent , macro culpture on frons rugulose , e pi torular sculpture rugulose. Four clypeal setae , medial longest , mandible unknown . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina weak, complete , medially present but indistinct . Distance between lateral ocellus and hyperoccipital carina approximately one ocellar diameter .

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64 Mesosoma. Epomial carina , present , complete or nearly so . Lateral pronotal microsculpture present throughout. Mesoscutellar spine long and blunt , pointing posteriorly in lateral view . Propodeal keel rounded in lateral view. Dorsal m esosoma weakly convex in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines weakly impressed . Seta e o n mesoscutum moderately dense. M esoscutal lamella short , rounded, narrow. Seta e o n mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S 2 present in narrow band at posterior margin. T2 smooth, with narrow band of microsculpture at posterior margin . T2 shorter than or equal to length of mesosoma. Forew ing. Setae on disc short , dense . D isc uniformly setose distally, proximally glabrous with linea setosa. S etal fringe uniform in length on posterior and anterior margins. Diagnosis. Important characters for identifying Synopeas sp. 10 include the weak hyperoccipital carina and the microsculpture of the lateral pronotum, which is absent only in the ventralmost portion . It can be distinguished from Synopeas sp. 13 by the pattern of microsculpture on S2. In Synopeas sp. 1 0, the microsculpture is limited to a narrow band along the posterior margin that is half as wide as that of Synopeas sp. 13 (Figure 2 25) , and Synopeas sp. 13 also has a well developed hyperoccipital carina. The notauli of Synopeas sp. 10 are not indicated, which allows it to be easily separated from Synopeas sp. 9 and Synopeas sp. 9b. Plant Associations. Reared from blister gall on Nauclea sp. (Rubiaceae ).

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65 Material E xamined . 1 female specimen . Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid gall on Nauclea sp. 1. Emerged 30 XI 2010. Figure 2 14 . Synopeas species 10 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 11 , incertae sedis Description. Body length 0.9 mm. Body brown , coxae brown, legs otherwise yellow . Head in fron tal view roughly circular . Frontal keel absent, microsculpture on frons reticulate, epi torular sculpture reticulate with minute rugulae. Four clypeal setae, medial

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66 setae longest , m andible bidentate . Ocular ocellar length approximately one ocellar diameter . Hyperoccipital carina absent . Mesosoma. Epomial carina present . Lateral pronotal microsculpture present dorsally, smooth ventrally. Mesoscutellar spine inconspicuous . Propodeal keel rounded in lateral view. Mesosoma dorsally convex in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines very faint. Setae sparse on mesoscutum. Mesoscutal lamella broad, square . Setae on mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S 2 present in narrow band at posterior margin . T2 as long as mesosoma. Forew ing. Unknown. Diagnosis. The head of this species is circular in anterior view , not distinctly wider than high . It lacks a hyperoccipital carina, and the scutel lar spine is inconspicuous or absent. The shape of the head sets it apart from Synopeas spp. 7 and 22, which have ovoid heads in frontal view , and from Synopeas sp. 6, which has a block y head in lateral view . It may be distinguished from Synopeas sp. 8 by the lateral pronotal microsculpture, which i n Synopeas sp. 11 is absent in the ventral third. Plant Associations. Reared from gall s on Saurauia poolei (Actinidaceae). Material Examined . 1 female specimen . Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid gall on Saurauia poolei. Emerged 01 VII 2010.

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67 Figure 2 15 . Synopeas species 11 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 13 , new species Description. Body length 1.2 1.6 mm. Body black , coxae brown, l eg s otherwise yellow to brown . Head in frontal view slightly ovoid. Central keel absent , microsc ulpture on frons reticulate , e pi torular sculpture reticulate with minute rugulae. Four clypeal setae , medial longest , evenly spaced, or medial setae closer to each other than to la teral setae ; mandible bidentate . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina robust , complete. Distance between lateral ocellus and hyperoccipital carina less than or equal to one ocellar diameter . Mesosoma. Epomial carina present , complete or nearly so . Lateral pronotal microsculpture present dorsally, smooth ventrally. Mesoscutellar spine long and blunt ,

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68 pointing posteriorly in lateral view . Propodeal keel pointed dorsally and ventrally in lateral view. Mesosoma do rs ally convex in lateral view . Notauli unmarked or faintly suggested. Parapsidal lines very faint. Seta e sparse o n mesoscutum. M esoscutal lamella lon g, narrow, pointed . Seta e o n mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S 2 present in wide band at posterior margin. T2 with reticulate microsculpture in posterolateral corners. T2 shorter than or equal to length of mesosoma. Forew ing. Setae on disc short , dense . D isc uniformly setose distally, proximally glabrous with linea set osa. S etal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. Synopeas sp. 13 is similar to Synopeas sp. 10, from which it can be separated by the pattern of microsculpture at the posterior margin o f S2 . In Synopeas sp. 13 , the sculpture forms a wide band, longer than S3 (Figure 2 25) , and in Synopeas sp. 10 it is limited to a very narrow band . The absent or faint notauli of Synopeas sp. 13 enable it to be easily separated from Synopeas spp. 9 and 9b. Synopeas sp. 13 has a dorsally convex and blunt mesoscutellar spine, distinguishing it from Synopeas spp. 1 and 1b, both of which have a short, pointed mesoscutellar spine. Plant Associations. Reared from fuzzy and nodule like galls on Cypholophus (Urticaceae). Material E xamined . 4 female specimen s. Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid galls on Cypholophus sp. 1 . 1 female emerged 28 III 2012. 1 female emerged 29 III 2012. 1 female emerged 10 VII 2012. 1 female emerged 17 VII 2012.

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69 Figure 2 16 . Synopeas species 13 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 15 (cf . Synopeas psychotriae Buhl) Description. Body length 1.3 1.4 mm. Body black to dark brown , coxae brown, l eg s otherwise yellow to brown . Head in frontal view circular to slightly ovoid. Central keel absent , impressed line extending from between toruli to median ocellus , micros culpture on frons reticulate , e pi toru l ar sculpture reticulate with minute rugulae. Four clypeal setae , medial longest , closer to each other than to lateral setae , mandible bidentate . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina weak, complete. Distance between lateral ocellus and hyperoccipital carina less than or equal to one ocellar diameter .

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70 Mesosoma. Epomial carina present , complete or nearly so . Lateral pronotal microsculpture present dorsally, smooth ventrally. Mesoscutellar spine short and pointed , angled posteriorly or posterodorsally in lateral view . Propodeal keel dorsally and ventrally pointed in lateral view. Mesosoma dors ally convex in lateral view. Notauli unmarked or faintly suggested. Parapsi dal lines well impressed . Seta e sparse o n mesoscutum. M esoscutal lamella broad and square. Seta e o n mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S 2 present in narrow band at posterior margin. T2 smooth , or with very narrow b and of microsculpture on posterior margin . T2 shorter than or equal to length of mesosoma. Forew ing. Setae on disc short , sparse . D isc uniformly setose distally, glabrous proximally. S etal fringe approximately uniform in length on posterior and anterior margins. Diagnosis. The parapsidal lines of Synopeas sp. 15 are well impressed , distinguishing it from the similar Synopeas sp.3, which has unmarked or very faint parapsidal lines. The metasomal microsculpture of Synopeas sp. 15 is minimal, being absent or restricted to very narrow bands at the posterior margins of each segment , whereas Synopeas spp. 1, 1b, and 13 have more extensive metasomal sculpturing (Figure 2 25) . The meso scutellar spine may be angled posteriorly or posterod orsally . Comments. The female holotype of S. psychotriae has a posterodorsally pointed mesoscutellar spine. The male Synopeas sp. 15 has a posteriorly pointed mesoscutellar spine, while that of the female from the same gall is posterodorsally pointed. Synopeas

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71 psychotriae and Synopeas sp. 15 are otherwise morphologically and ecologically similar. The angle of the mesoscutellar spine may be sexually dimorphic. Plant Associati ons. Reared from galls on Psychotria ramuensis (Rubiaceae). Material E xamined . 1 female, 1 male. Papua New Guinea, Morobe Province, Ohu , 5.233 °S , 14 5.686 °E , Coll. P. Butterill. Ex. cecidomyiid gall on Psychotria ramuensis . Emerged 22 II 2011. F igure 2 17 . Synopeas species 15 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad.

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72 Synopeas sp. 16 (cf . Synopeas butterilli Buhl) Description. Body length 1.3 mm. Body black , coxae brown, l eg s otherwise yellow to brown . Head in frontal view circular . Central keel complete, very weak, may be difficult to see ; micros culpture on frons reticulate , e pi torular sculpture rugulose. Four clypeal setae , medial longest , evenly spaced, or medial setae somewhat closer to each other than to lateral setae ; mandible bidentate . Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina robust , complete. Distance between lateral ocellus and hyperoccipital carina greater than one ocellar diameter . Mesosoma. Epomial carina present, complete or nearly so. Lateral pronotal microsculpture present in narrow line medially , smooth ventrally and dorsally. Mesoscutellar spine short and pointed , pointing posteriorly or posterodorsally in lateral vi ew. Propodeal keel rounded in lateral view. Mesosoma dors ally convex in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines very faint. Seta e sparse on mesoscutum. M esoscutal lamella short and rounded , projecting less than its width beyon d margin. Seta e on mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S 2 present in narrow band at posterior margin. T2 with narrow band of microsculpture at posterior margin . T2 shorter than or equal to length of mesosoma. Forew ing. Setae on disc short , sparse . D isc setose distally, proximally glabrous with linea setosa. S etal fringe approximately uniform in length on posterior and anterior margins.

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73 Diagnosis. The lateral ocellus and the hyperoccipital carina are widely separated in Synopeas sp. 16, by approximately 1.5 ocellar diameters. The sculpture of the lateral pronotum is restricted to a narrow line medially, and the lateral pronotum is smooth dorsally and ventrally. These two characters set it apart from ot her species with a short, pointed mesoscutellar spine: Synopeas spp. 1 , 1b, 2, 3, and 4. Comments. The female holotype of S. butterilli has a posterodorsally pointed mesoscutellar spine. The male Synopeas sp. 16 has a posteriorly pointed mesoscutellar spin e. Synopeas butterilli and Synopeas sp. 16 are otherwise morphologically and ecologically similar. The angle of the mesoscutellar spine may be sexually dimorphic. Plant Associations. Reared from galls on Paramyristica cf. sepicana ( Myristicaceae). Occasionally found in the same gall as Inostemma sp. Material E xamined . 1 male specimen . Papua New Guinea, M adang Province, Mis . 5.183 °S , 14 5.758 °E , Coll. P. Butterill. Ex. cecidomyiid gall on Paramyristica cf. sepicana . Emerged 16 VII 2010 .

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74 Figure 2 18 . Synopeas species 16 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 19 , new species Description. Body length 1.2 mm. Body dark brown , coxae brown, l eg s yellow to brown. Head in frontal view ovoid. Centra l keel weak, complete ; frons with reticulate microsculpture medially, rugose macrosculpture dorsally ; e pi torular sculpture rugulose. Four clypeal setae , medial longest , closer to each other than to lateral setae , mandible bidentate . Ocular ocellar length approximately one ocellar diameter . Hyperoccipital carina absent. Mesosoma. Epomial carina present, complete or nearly so . Lateral pronotal microsculpture present dorsa lly, smooth ventrally, with smooth band anterior to tegula.

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75 Mesoscutellar spine inconspicuous. Propodeal keel rounded in lateral view. Dorsal m esosoma weakly convex in lateral view. Notauli distinctly grooved . Parapsidal lines very faint. Seta e sparse o n m esoscutum. M esoscutal lamella truncate, with striate sculpture. M esoscutellum striate ly sculpture d, laterally setose . Metasoma. S2 and T2 smooth . T2 shorter than mesosoma. Forew ing. Setae on disc medium in length, moderately dense . D isc uniformly setose di stally, proximally glabrous with broad linea setosa. S etal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. The head of Synopeas sp. 19 has reticulate microsculpture on the frons , with rugose macrosculpture extending from the median ocellus to the occiput. The notauli are distinctly grooved . Synopeas spp. 9 and 9b also have rugose head sculpture and grooved notauli, but both species have a prominent mesoscutellar spine, whereas in Synopeas sp. 19 the mesoscutellar spine is absent or inconspicuous . Plant Associations. Reared from blister galls on Nauclea sp. (Rubiaceae). Found in the same gall as Synopeas sp. 4. Material E xamined . 1 male specimen . Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid gall on Nauclea sp. 1. Emerged 26 XI 2010.

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76 Figure 2 19 . Synopeas species 19 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 22 , new species Description. Body length 1.1 1.2 mm. Body black , coxae brown, l eg s otherwise dark yellow to brown. Head in frontal view distinctly ovoid. Central keel absent , micros culpture on frons reticulate , e pi torular sculpture reticulate with minute rugulae. Medial clypeal setae widely spaced , about as far apart as toruli ; mandible unidentate. Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina absent, vertex angled sharply onto occiput, posterior vertex medially depressed . Distance between lateral ocellus and hyperoccipital carina greater than one ocellar dia meter .

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77 Mesosoma. Epomial carina present, complete or nearly so, posteriorly somewhat weakened. Lateral pronotal microsculpture present dorsally, smooth ventrally. Mesoscutellar spine inconspicuous. Propodeal keel pointed dorsally and ventrally in lateral view. Dorsal m esosoma strongly convex in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines weakly impressed . Seta e sparse o n mesoscutum. M esoscutal lamella triangular, posterior edges defined by two carinae , not projecting much beyond posterior margin of mesoscutum . Seta e o n mesoscutellum laterally dense, medially sparse. Metasoma. Microsculpture on S 2 present in narrow band at posterior margin. T2 smooth , or with very narrow band of microsculpture at poster ior margin. T2 shorter than mesosoma. Forew ing. Setae on disc moderately long , moderately dense . D isc uniformly setose distally, proximally glabrous with linea setosa. S etal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. T he posterior vertex of Synopeas sp. 22 is medially depressed, which is unique to the species treated here. In lateral view, Synopeas sp. 22 has an extremely convex mesosoma and a wide pronotum , giving it a hunchbacked appearance. This distinguishes it from Synopeas sp. 7, which has a similarly shaped head. Synopeas sp. 22 resemble s the description of S. quasimodo Buhl , an Indonesian species. However, examination of the holotype of S. quasimodo reveals only a modestly convex mesosoma, as well as a completely sculptured lateral pronotum and a strong, complete hyperoccipital carina (Figure 2 30) . Plant Associations. Reared from galls on Nauclea sp. (Rubiaceae).

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78 Material E xamined . 1 male specimen . P apua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid gall on Nauclea sp. 1 . Emerged 26 XI 2012. Figure 2 20 . Synopeas species 22 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 23 , new species Description. Body length 1 mm . Body dark brown , coxae brown, l eg s otherwise yellow to brown. Head in frontal view ovoid. Central keel present ventrally, absent medially, may be present just ventral to median ocellus ; micros culpture on frons reticulate, with some minute rugulae around median ocellus ; e pi torular sculpture rugose. Clypeal seta e

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79 unknown, mandible unknown. Ocular ocellar length less than one ocellar diameter . Hyperoccipital carina absent. Mesosoma. Epomial carina present , complete or nearly so . Lateral pronotal microsculpture present dorsally, smooth ventrally. Mesoscutellar spi ne short and pointed , angled posterodorsally in lateral view . Propodeal keel pointed dorsally and ventrally in lateral view. Mesosoma dors ally flattened in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines very faint or marked by absenc e of sculpture. Seta e sparse o n mesoscutum. M esoscutal lamella longer than wide , with rounded tip . M esoscutellum medially glabrous , laterally setose . Metasoma. S2 smooth. T2 sculpture unknown. T2 shorter than mesosoma. Forew ing. Setae on disc medium in length, moderately dense . S etal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. Synopeas sp. 23 may be distinguished from other species with a short, pointed mesoscutellar spine by the absence of a hyperoccipital carina an d the flattened appearance of the mesosoma in lateral view. Plant Associations. Reared from blister galls on Nauclea sp. (Rubiaceae). Material E xamined . 1 male specimen. Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid gall on Nauclea sp. 1 . Emerged 24 VIII 2010.

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80 Figure 2 21 . Synopeas species 23 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas sp. 24 , new species Description. Body length 0.9 mm. Body black to dark brown , coxae brown, l eg s otherwise yellow to brown. Head in frontal view slightly ovoid. Central keel absent , micros culpture on frons reticulate , e pi torular sculpture rugulose. Clypeal setae unknown, mandible bidentate . Ocular ocellar length approximately one ocellar diameter . Hyperoccipital carina complete, medially weakened . Distance between lateral ocellus and hyperoccipital carina approximately one ocellar diameter .

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81 Mesosoma . Epomial carina present, complete or nearly so . Lateral pronotal microsculpture present dorsally, smooth ventrally. Mesoscutellar spine short and pointed , pointing posteriorly in lateral view . Propodeal keel rounded in lateral view. Mesosoma dors ally conv ex in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines very faint. Seta e sparse o n mesoscutum. M esoscutal lamella long and narrow , with rounded tip . Seta e o n mesoscutellum laterally dense, medially sparse. Metasoma. S2 smooth . T2 smooth , or with very narrow band of microsculpture at posterior margin . T2 shorter than or equal to length of mesosoma. Forew ing. Setae o n disc medium in length, moderately dense . S etal fringe distinctly longer on posterior margin than on anterior margin. Diagnosis. Identification of Synopeas sp. 24 requires the use of numerous characters because it does not have any obviously distinct features . It is most similar to Synopeas sp. 2, from which it can be separated by the development of the hyperoccipital c arina, which is medially weakened in Synopeas sp. 24 and uniformly robust in Synopeas sp. 2. It is also similar to Synopeas sp. 3, but Synopeas sp. 24 has no central keel, while Synopeas sp. 3 has a partial central keel. Plant Associations. Reared from red galls on Nauclea sp. (Rubiaceae). Material E xamined . 1 female specimen. Papua New Guinea, Morobe Province, Yawan , 6.1414142 °S , 146.8741517 °E , Coll. P. Butterill. Ex. cecidomyiid gall on Nauclea sp. 1 . Emerged 25 XI 2010.

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82 Figure 2 22 . Synopeas spe cies 24 . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad. Synopeas pterocarpi Buhl Description. Body length 1.4 1.5 mm . Body black, coxae brown, legs otherwise yellow to brown . Head in frontal view circular . Central keel absent, micros culpture on frons reticulate , e pi torular area sculpture d with parallel, arched rugae. Four clypeal setae , medial longest , evenly spaced, mandible unknown . Ocular ocellar length approximately

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83 one ocellar diameter . Hyperoccipital carina robust , complete. Distance between lateral ocellus and hyperoccipital carina approximately one ocellar diameter. Mesosoma. Epomial carina present, complete or nearly so. Lateral pronotal microsculpt ure present dorsally, smooth ventrally. Mesoscutellar spine inconspicuous or absent. Propodeal keel rounded in lateral view. Mesosoma dors ally convex in lateral view. Notauli unmarked or faintly suggested. Parapsidal lines very faint. Seta e sparse o n mesos cutum. M esoscutal lamella broad and roun ded. Setae on m esoscutellum medially sparse , laterally den se . Metasoma. Microsculpture on S 2 unknown. T2 smooth . T2 shorter than or equal to length of mesosoma. Forew ing. Setae on disc short, sparse, a rrangement unknown. Diagnosis. The frons of S. pterocarpi is characterized by parallel, arched rugae above the toruli, not found in other Synopeas from New Guinea. Plant Associations. Reared from galls on Pterocarpus indicus (Fabaceae). Material E xamined. Morphology of holotype examined from images taken by Elijah Talamas.

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84 Figure 2 23 . Holotype of Synopeas pterocarpi . Clockwise from top left: lateral habitus, anterior head , posterodorsal habitus, dorsal habitus. Scale bar represents 0. 1 mm. Photos by Eli jah Talamas. Synopeas c s oszi Buhl Description . Body length 1.7 2.3 mm. Body reddish brown, coxae brown, legs otherwise yellow to brown. Head in frontal view ovoid . Central keel absent, microsculpture on frons reticulate, epi torular sculpture rugulose . Clypeal setae unknown, mandible toothed. Ocular ocellar length approximately one ocellar length. Hyperoccipital carina strong, complete. Distance between lateral ocellus and hyperoccipital carina approximately one ocellar diameter .

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85 Mesosoma. Epomial carina present, complete . Lateral pronotal microsculpture present anterodorsally, absent posteroventrally. Mesoscutellar spine long and thin, with rounded tip, pointing posteriorly in lateral view. Mesosoma dorsally convex in lateral view . Notauli unmarked or faintly suggested. Parapsidal lines very faint. Seta e o n mesoscutum medially sparse, laterally dense. Mesoscutal lamella broad and rounded or roughly triangular. Seta e o n mesoscutellum medially sparse, laterally dense. Metasoma. Microsculpture on S 2 present in narrow band at posterior margin. T2 smooth with narrow band of microsculpture at posterior margin. T2 extremely elongate, longer than head and mesosoma combined. Forew ing. Setae on disc short, dense . D isc uniformly setose distally, proximally glabrous with broad linea setosa. S etal fringe approximately uniform in length on posterior and anterior margins. Diagnosis. This species is easily recognized by the extraordinary length of the second metasomal segment , which is longer than the head and mesosoma combined. The metasoma is narrow anteriorly, expanding posteriorly to become wider than the rest of the body at the junction between metasomal segments 2 and 3. The shape of the anten nal scape is unusually expanded distally forming a club, then contracting into a narrowly curved apex (Figure 2 24 ). Plant Associations. Unknown . Material E xamined. Morphology of holotype examined from images taken by Elijah Talamas.

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86 Fig ure 2 24 . Holotype of Synopeas csoszi . Scale bar represents 0. 5 mm. Photo by Elijah Talamas. Key to the Species of Synopeas of New Guinea 1a. T2 longer than the head and mesosoma combined (Figure 2 24 ... .................................................................................................. Synopeas csoszi Buhl 1b. T 2 equal to or less than length of mesoso ma (Figures 2 3, 2 4) . ................ 2 2a. Notaul us distinctly grooved (Figures 2 12, 2 19) .. ............ 3 2b. Notaul us unmarked or faintly suggested (Figures 2 20, 2 21) .................................. 5

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87 3a. Mesoscutellar spin e absent or inconspicuous (Figure 2 19) ; lateral portion of frons without r ugose macrosculpture ... ... ....................................................... Synopeas sp . 19 3b. Mesoscutellar spine distinct and conspicuous ; lateral portion of frons with r ugose macrosculpture ... ... ......................................................................................................... 4 4a. Length of m esos utellar spine only slightly greater than its thickness in lateral view; lateral pronotum with microsculpture dorsally, smooth ventrally, with smooth band anterior to tegula (Figure 2 12) ...... ......................................................... Synopeas sp . 9 4b. Length of mesoscutellar spine more than twice its thickness at i ts midpoint; l ateral pronotum with microsculpture dorsally, smooth ventrally (Figure 2 13) ......................... .............................................................................................................. Synopeas sp . 9b 5a. Epomial carina absent or reduced, extending less than halfway up pronotum ......... ............................................................................................................... ... Synopeas sp . 6 5b. Epomial carina present , extending more than halfway up pronotum ...... .................. 6 6a. Mesoscutellar spine absent or inconspicuous (Figure 2 30, top row) ... ................. .. . 7 6b. Mesoscutellar spine long or pointed (Figure 2 30, center and bottom row ) .. ... ....... 12

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88 7a. Head large and triangular in anterior view; dorsal margin of c ompound eyes co planar with vertex ; length of malar space about equal to height of compound eye.. ...... .......................................................................................... ............... .... ..... Synopeas sp . 5 7b. Head round or ovoid in anterior view .. ...................................................................... 8 8a. Hyperoccipital carina present and complete ... ................................................ .......... 9 8b. Hyperoccipital carina absent , very weak , or incomplete ... ...................................... 1 0 9a. Frons with parallel, arched rugae above torulus; l ateral pronotum with microsculpture absent in ventral third (Figure 2 23) ... ..... ..... Synopeas pterocarpi Buhl 9b. Frons without parallel, arched rugae (but may have minute, irregular epiclypeal rugulae); l ateral pronotum with microsculpture throughout or with narrow smooth area at ventral apex (Figure 2 11) ... ................................................................... Synopeas sp. 8 10a. Head in frontal view circular , not distinctly wider than high... ........ Synopeas sp . 11 10b. Head in frontal view ovoid, distinctly wider than high.. ........................................... 11 11a. Mesosomal dorsum strongly convex in lateral view (Figure 2 20 ) ; v ertex medially depressed .. ....................................................... ..................................... Synopeas sp . 22 11b. Mesosomal dorsum moderately convex in lateral view (Figure 2 10 ); v ertex rounded onto occiput.. ............................................................................. Synopeas sp . 7

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89 12a. M esoscutellar spine long and blunt (Figures 2 14, 2 16) ...................................... 13 12b. Mesoscutellar spine short and pointed (Figure 2 30, center row) ......................... 14 13a. Hyperoccipital carina weak ly developed and incomplete ............. Synopeas sp . 10 13b. Hyperoccipital carina robust and complete ................................... Synopeas sp . 13 14a. Central keel well developed and extending to median ocellus (Figure 2 7 ) .............. ................................................................................................................ Synopeas sp . 4 14b. Central keel absent, weakly developed, or not extending to median ocellus (Figures 2 6, 2 18, 2 22 ) . ..................... ........................................................................ . 15 15a. Hyperoccipital carina absent ........................................................ Synopeas sp . 23 15b. Hyperoccipital carina present , carina may be weak or incomplet e ....................... 16 16a. Patch of microsculpture on posterior S2 relatively wide , longer than S3 .............. 17 16b. S2 with no microsculpture or with very narrow band at posterior margin ............. 18 17a. T 2 with wide (longer than T3) band of microsculpture at posterior margin ................ ................................................................................................................. Synopeas sp . 1 17b. T2 without microsculpture , or with very nar row band at posterior margin .................. .............................................................................................................. Synopeas sp . 1b

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90 18a. Parapsidal lines well impressed (Figure 2 17) ; OOL less than 1 ocellar diam eter .... ............................................................................................................... Synopeas sp . 15 18b. Parapsidal lines unmarked or faintly indicated (Figure 2 18) ; OOL variable ........ 19 19a. Dorsal margin of l ateral pronotum smoot h ................................... Synopeas sp . 16 19b. Dorsal margin of lateral pronotum with microsculpture ......................................... 20 20a. Hyperoccipital carina robust and complete ..................................... Synopeas sp . 2 20b. Hyperoccipital carina weak or incomplete ............................................................ 21 21a. Central keel partial, present in ventral but not dorsal part of frons; forewing with short fringe of setae approximately uniform in length on anterior and posterior margins (Figure 2 27, top left) .............................................................................. Synopeas sp. 3 21b. Central keel absent; forewing with fringe of setae noticeably longer on posterior margin than on anterior margin (Figure 2 27, top right) ........................ Synopeas sp. 24 Discussion The condition of the specimens from the Noona Dan is rather poor. Many are pale yellow , which is not typical for Synopeas and suggests that they may have b een bleached by the sun or chemical processes. The cuticle of the se specimens is translucent, rendering visible the shriveled musculature and making it difficult to discern microsculpture. Of course, it is challenging to keep M alaise trap samples in good condition on a boat in the tropics for two years . The scientists and crew of the Noona Dan experienced many political and medical difficulties , including the death of

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91 ornithologist Erik Petersen , who was reportedly killed by an unspecified venomous arthropod (Wolff, 1963) . Keeping alcohol samples out of the sun may have been a relatively low priority for much of the journey. Though the causes ar e understandable , the poor quality of these samples is a n impediment to interpreting their morphology . Should fresh representatives of these species be collected, it may be difficult to match them to the deteriorated holotypes. Th e discernable morphology of the Noona Dan material did not match that of the Butterill or B alogh material , but it is important to note the harmful result of describing species from damaged specimens. For this reason, Synopeas sp. 11 from my study is designated incertae sedis . Morp hospecies 11 has a thin, pale cuticle, and did not yield any barcode data , indicating that the material is damaged . Scanning electron microscopy may be necessary to circumvent the limitations of light microscopy. The results of this study emphasize the imp ortance of certain characters to the morphological identification of Synopeas species. P atterns of cuticular microsculpture on the lateral pronotum and metasoma proved especially useful for separating species, as well as patterns of macrosculpture on the h ead. The ocular ocellar length (OOL) was found to be a useful morphometric character in some instances . The proportions of the metasoma can also aid in species identification, although it is possible that this character is only applicable to female specimens. The arrangement and lengths of the clypeal setae can be useful for separating species . However, the clypeal setae cannot always be seen without removing the antennae or remounting the specimens. All Synopeas specimens examined in this study had four clypeal setae, and this may be stable at the generic level.

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92 The meso scutellar spine is among the mo re obvious characters available for identification of Synopeas , being visible at lower magnifications than cuticular sculpture and setal patterns. In so me cases, such as the distinction between Synopeas spp. 9 and 9b, subtle differences in spine shape were informative. In the cases of Synopeas spp. 15 and 16, tentatively identified as S. psychotriae Buhl and S. butterilli Buhl , respectively, the utility o f the spine shape i s questionable. The s pecimen s examined for this thesis differed from the type material in shape and angle of meso scutellar spine but were otherwise morphologically and ecologically similar. In the absence of barcode data from name bearing specimens, there was in sufficient evidence for proposing new species. It is worth noting that male specimens had the spine pointing posteriorly, while female specime ns had the spine pointing posterodorsally. Such variation may be sexually dimorphic, although it is also possible that th e characte r is variable in one or both sexes. The character of meso scutellar spine angle should be approached with caution. Synopeas sp . 5 exhibits a morphological oddity in the shape, size, and proportions of the head. It is interesting to note that an Inostemma species with similar head morphology (Figure 2 28 ) emerged from the same host plant ( Macaranga ). Other Macaranga reared Synopea s have a more typical head. There is also a Platygaster species with an exceptionally large head described from New Guinea, but its host associations are unknown. It would be interesting to learn if this species also inhabits galls on Macaranga. Th e specie s, based on two female specimen s , w as proposed to represent a new subgenus, Rhynchoplatygaster Buhl ( Buhl, 2013 ) .

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93 This striking example of convergence suggest s several possible ecological interpretations. The large head may house powerful mandibular muscles for chewing an exit hole out of a hard gall, or for inter or intraspecific combat. The wide malar space may also indicate the presence of enlarged cephalic glands of unknown function and significance. In any case, the large triangular head is probably ecologically adap tive and is not indicative of a separate lineage. Due to the severity of taxonomic disarray in Synopeas , it is currently impossible to compare a given specimen to all other described species of Synopeas worldwide. Without keys or thorough descriptions, c omparing putative species to the known fauna of the region is as much as a student can do within a few years. This presents a dilemma to the aspi ring hymenopterist: should one propose new names for the described species , despite the chance that they are also found on distant continents , or sufferer of the mihi itch (Evenhuis , 2008) will surely affix his own name to the species at the first opportunity ? There is no easy answer to this questio n . Having ecological context for the putative species can provide some idea of their possible geographic distribution. If the gall forme rs are specialists, as is gener ally the case, and the gall host plants are restricted to New Guinea or to Australasia, then it is more likely that their parasitoids are also restricted to this region (Dorchin et al. , 2019; Miller and Raman , 2019) . If the p lants have a wide geographic distribution, it may be that the gall formers and their parasitoids do as well. As it happens, the plant specie s from the Butterill project have a limited geographic distribution (Table 2 4 ; P lants of the World Online , 2019; Le enhouts, 1960 ) . Some plants could not be identified to species ,

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94 suggesting that they are rare, understudied, or endemic . Therefore, I conclude that the Synopeas species described in my thesis are not likely to occur outside of Australasia , and that comparison to type material of known fauna of the region is sufficient grounds for the proposal of new species names . The relationship between morphospecies, COI barcode data, and gall host plant identi ty is explored further in Chapter 3. Table 2 4 . Known geographic distributions of gall host plants for Synopeas specimens from the Butterill loan to the Florida State Collection of Arthropods . Host plant family Host plant species Known distribution Actinidaceae Saurauia conferta Papua New Guinea , Solomon Islands Saurauia poolei Papua New Guinea Saurauia schumanniana Papua New Guinea , Solomon Islands Elaeocarpaceae Elaeocarpus sp. 1 Unknown Euphorbiaceae Homalanthus nervosus Papua New Guinea Macaranga strigos a Papua New Guinea Loganiaceae Neuburgia corynocarpa Papua New Guinea, Solomon Islands , Fiji Meliaceae Aglaia rimosa Papua New Guinea, Indonesia, Philippines, Taiwan Myristicaceae Paramyristica cf. sepicana Papua New Guinea Piperaceae Piper amboinense Indonesia Piper celtidiforme Papua New Guinea, Indonesia, Philippines Rubiaceae Nauclea sp. 1 Unknown Nauclea tenuiflora Papua New Guinea Psychotria ramuensis Papua New Guinea Urticaceae Cypholophus sp. 1 Unknown Cypholophus friesianus Papua New Guinea

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95 Figure 2 2 5 . Comparative morphology of the second metasomal sternite (S2). Clockwise from top left: Synopeas spp. 1, 1b, 2, 3, 6, and 13. Photos by Jessica Awad.

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96 Figure 2 2 6 . Comparative morphology of the second metasomal tergite (T2). Clockwise from top left: Synopeas spp. 1, 1b, 13, and 9. Photos by Jessica Awad.

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97 Figure 2 2 7 . Comparative morphology of the forewing. Clockwise from top left: Short and sparse setae with fringe unifor m in length ( Synopeas sp. 16) ; Long and sparse setae with fringe longer on posterior margin ( Synopeas sp. 24) ; Short and dense setae, proximally glabrous with linea setosa ( Synopeas sp. 2) ; Short and dense setae with fringe longer on posterior margin ( Synopeas sp. 13) . Photos by Jessica Awad.

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98 Fi gure 2 28 . Inostemma specimen reared from cecidomyiid gall on Macaranga . Clockwise from top left: lateral habitus, ventral habitus, dorsal habitus. Scale bar represents 0.2 mm. Photos by Jessica Awad.

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99 Figure 2 29 . Holotype of Synopeas quasimodo . Clockwise from top left: lateral habitus, ventral h abitus , dorsal habitus. Photos by Elijah Talamas.

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100 Figure 2 30 . Comparative morphology of the mesoscutellar spine. Top row: absent or inconspicuous. Center row: short and pointed. Bottom row: long and blunt. Photos by Jessica Awad.

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101 CHAPTER 3 PHYLOGENETIC ANALYSIS Introduction The development of a robust barcode library is essential to molecular identification. Much work remains to be done to further the utility of this important tool for diagnosis of Synopeas species . For example, a search of the BOLD Systems online repository (Ratnasingham and Hebert, 2007) for the term Synopeas 2,2 62 sequences as of April 2020 . Only 399 (17.6%) records had species names , and these records collectively represented only 4 species. The sequences formed 136 BINs or clusters, suggesting that the database includes 132 species that have not been identifie d (Ratnasingham and Hebert, 2013) . Thus, the utility of these sequences for Synopeas species identification is currently very limited. The phy logenetic analysis of COI barcode data provides structure and context to species descriptions. While it i s important to relate barcode sequences to morphological and taxonomic informatio n for diagnostic purposes , arranging these sequences in a tree can lead to further insight. Ph ylogenetic analysis reveals patterns in the relationships amo n g morphology, ecolo gy, geography, and genetics. In a relatively large genus such as Synopeas , phylogenetics can help define species groups, breaking down the taxon into more manageable pieces for detailed attention in the future . Maximum likelihood was chosen as the most appropriate optimality criterion for these data. Evolutionary models of invertebrate mitochondrial DNA have been well developed ( Pen tinsaari et al. , 2016 ), so a model based method is potentially more informative than a parsimony analysis. However, in the name of thoroughness, I also

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102 performed a parsimony analysis and compared the results , an approach referred to by Wheeler (2012) as th . Bayesian techniques were considered unnecessary for a small, single gene data set. Materials and Methods Material Examined Between one and seven specimens were selected f or DNA extraction from each morphospecies of Synopeas desc ribed in Chapter 2 . Selection criteria included number available, size variation, morphological variation, and gall host plant identity. Six specimens of Inostemma and seven specimens of Platygaster , al so reared in New Guinea by Philip Butterill, were sele cted as outgroups. For geographic reference, DNA was extracted from three Synopeas specimens from the USA, the UK, and Myanmar. Synopeas holotypes from the Butterill collection , along with the Myanmar Synopeas specimen, are to be deposited in the United States National Museum of Natural History in Washington, DC. Synopeas paratypes , other Platygastrinae from the Butterill collection , and the US Synopeas specimen will be deposited in the Florida State Collection of Arthro pods in Gainesville, Florida. The Synopeas specimen from the UK will be deposited in the State Museum of Natural History in Stuttgart, Germany, where it can be included in future studies o f the Synopeas of Europe. Molecular Techniques DNA extraction used a non destructive protocol. The protocol was adapted for microhymenopterans by John Noyes, John Heraty, and Hua yan Chen and is a

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103 modification of procedures from the QIAGEN DNeasy® Tissue Handbook (2006). Each specimen was softened in 70% ethanol for at l east 2 hours. Alcohol was removed and replaced with a digestion mix of 180 µl buffer ATL and 40 µL proteinase K. The specimen was incubated in the digestion mix at 55°C for 8 24 hours, then frozen at 20°C for another 8 24 hours. The sample was thawed at r oom temperature, and the supernatant was separated from the specimen. DNA was extracted from the supernatant by using the buffers and collection tubes provided in the DNeasy® blood and tissue kit (QIAGEN Group, Hilden, Germany). When the supernatant was re moved from the specimen tube, it was replaced with 300 µL distilled water. The specimen remained in water for 45 minutes, after which 700 µL 95% ethanol was added. A specimen may be stored in this solution indefinitely. For morphological analysis, the spec imens were removed from alcohol and quickly dried upon a porous ceramic plate (Soilmoisture Equipment Corp., Goleta, CA). Once dry, the specimens were point mounted and imaged. DNA extracts were quantified with a NanoDrop 2000 spectrophotometer (Thermo Fis COI barcode region was amplified using primers LCO1490 and HCO2198. PCRs were conducted at 25 µL volumes with HiFi HotStart DNA Polymerase (Kapa Biosystems, Wilmington, MA). PCR products were verified by gel electrophor esis, then cleaned for sequencing by using the QIAquick Gel Extraction Kit (QIAGEN Group, Hilden, Germany). Purified PCR products were Sanger sequenced by using BigDye Terminator v3.1 on a SeqStudio Genetic Analyzer (Applied Biosystems, Thermo Fisher Scien tific, Waltham, MA). Sequencher 5.4.6 (Gene Codes

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104 Corporation, Ann Arbor, MI) was used for trimming sequence reads and assembling of sequence contigs. Analysis Barcode gene sequences were aligned using MUSCLE (Edgar, 2004) in the program MEGA version 7 ( Kumar et al. , 20 16 ) . In MEGA 7, the aligned sequences were used to generate a neighbor joining tree from 1 , 000 bootstrap replicates , using the Kimura 2 parameter method (Kimura, 1980) . The aligned sequences were evaluated in IQ TREE (Nguyen et al. , 2015) to select the best substitution model, which was GTR+F+R3 based on Akaike Information Criterion . The data were then analyzed in IQ TREE as a maximum likelihood tree reconstruction and no n parametric bootstrap with 1 , 000 replicates. Some Inostemma and Platygaster sequences were removed from this analysis, as they produced very long branches that made the tree unwieldy. The maximum likelihood analysis included 57 sequences , of which 50 were from Synopeas . Aligned sequences were analyzed in MEGA 7 using the maximum parsimony method , producing a bootstrap consensus tree from 1 , 000 replicates. Th e ana lysis used the Tree Bisection Regrafting (TBR) algorithm with search level 1, beginning with 10 initial trees formed by random addition of sequences. All codon positions were included, and positions with less than 95% coverage were eliminated. The maximum parsimony analysis included 63 sequences, of which 50 were from Synopeas . C OI sequences were compared to morphotypes to determine if morphologically similar specimens were also genetically similar. If high COI divergence was found in

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105 morphologically similar specimens, morphological and ecological data were re examined. Results A ll three analyses support ed the monophyly of the genus Synopeas (Figure 3 1, Figure 3 2, Figure 3 3) . Where multiple individuals from a morphospecies could be sampled, the COI sequences exhibited intraspecific differences of less than 3% (Figure 3 1) . In the maximum likelihood analysis (Figure 3 2) , each morphospecies was monophyletic . Most morphospecies had strong bootstrap support ( 100%) , except for Synopeas sp. 7 (86%) and Synopeas sp. 13 ( 89%) . Some clades were strongly supported. Species 1, 1b, 13 , and the specimen from the UK formed a clade with 99% support. Species 6 and 8 formed a clade with 98% support. A large and well supported clade (98%) included species 3, 4, 7, 15, 16, 22, 23, and 24. In the maximum parsimony analysis (Figure 3 3) , e ach m orphospecies was monophyletic , with strong bootstrap support (100%). Specimen PL91, originally designated as species 1, was later designated as Synopeas sp. 1b, based on morphological, ecological, and molecular data. Species 1, 1b, 13, and the specimen from the UK formed a clade with 96% support. A large and well supported clade ( 99%) included species 3, 4, 7, 15, 16, 22, 23, and 24. The New Guinea material did not all cl uster together. Synopeas specimens from the US, the UK, and Myanmar were placed relatively far apart from one another on the tree. In both maximum likelihood and parsimony analyses , the US and Myanmar specimens clustered with species 9, 9b, and 19 . Bootstr ap support was poor: 54% for

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106 maximum likelihood and 46% for parsimony. As stated above, the UK specimen clustered with species 1, 1b, and 13, with strong support in both analyses , although it was distinct from the New Guinea species . The specimen from the UK was tentatively identified as Synopeas larides (Walker) by comparison to the holotype photographed by Elijah Talamas. In general, each morphospecies cluster was associated with a single genus of plant (Table 3 1) . Synopeas sp. 1 was reared from Saurauia conferta and Saurauia schumanniana (Actinidaceae) . Synopeas sp. 2 was reared from Piper amboinense and P. celtidiforme (Piperaceae). Synopeas sp p . 4, 6, 8, and 9 were reared from galls on Nauclea (Rubiaceae). Synopeas sp p . 1b and 13 w ere reared fr om Cypholophus spp. (Urticaceae). Only Synopeas sp. 7 was associated with multiple host plant genera, having been reared from both Nauclea spp. and Macaranga strigosa (Euphorbiaceae). All other morphospecies in the analysis were represented by a single spe cimen.

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107 Table 3 1 . Gall host plant associations of Synopeas morpho species sampled for COI barcode analysis . Morphospecies Host plant species Number of specimens Synopeas sp. 1 Saurauia conferta 1 Saurauia schumanniana 2 Synopeas sp. 1b Cypholophus friesianus 1 Synopeas sp. 2 Piper amboinense 2 Piper celtidiforme 4 Synopeas sp. 3 Neuburgia corynocarpa 1 Synopeas sp. 4 N auclea sp. 1 7 Synopeas sp. 6 Nauclea sp. 1 1 Nauclea tenuiflora 1 Synopeas sp. 7 Nauclea sp. 1 2 Nauclea tenuiflora 2 Macaranga strigos a 2 Synopeas sp. 8 Nauclea sp. 1 2 Synopeas sp. 9 Nauclea sp. 1 5 Synopeas sp. 9b Nauclea sp.1 1 Synopeas sp. 13 Cypholophus sp. 1 4 Synopeas sp. 15 Psychotria ramuensis 1 Synopeas sp. 16 Paramyristica cf. sepicana 1 Synopeas sp. 19 Nauclea sp. 1 1 Synopeas sp. 22 Nauclea sp. 1 1 Synopeas sp. 23 Nauclea sp. 1 1 Synopeas sp. 24 Nauclea sp. 1 1

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108 Figure 3 1 . Neighbor joining tree of COI barcode data from Synopeas of New Guinea, including Platygaster and Inostemma species as outgroups, and Synopeas specimens from the UK, the USA, and Myanmar.

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109 Figure 3 2 . Maximum likelihood analysis of COI barcode data from Synopeas of New Guinea , including Platygaster and Inostemma as outgroups, and Synopeas specimens from the UK, the USA, and Myanmar .

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110 Figure 3 3 . Maximum parsimony bootstrap consensus tree of COI barcode data from Synopeas of New Guinea, including Platygaster and Inostemma as outgroups, and Synopeas specimens from the UK, the USA, and Myanmar .

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111 Discussion In general, different optimality criteria generated similar phylogen ies. All analyses agreed on the monophyly of Synopeas , of most clades, and of all morphospecies. The main differences were in the bootstrap values . For example, parsimony generated 100% support for all morphospecies clusters, while likelihood generated lower bootstrap support for Synopeas sp. 7 ( 92 %) and Synopeas sp. 13 ( 75 %) . Likelihood generated strong support (98%) for the monophlyly of Synopeas sp. 6 and Synop eas sp. 8 . Parsimony retrieved the same clade with only 86% support. However, the differences are rather minor . Combined with morphological and ecological data, the COI results lend support to the hypothesized morphospecies, although Synopeas spp. 7 and 13 should be carefully scrutinized . Some morphological characters formed discernible phylogenetic patterns. T he clade including Synopeas spp. 9, 9b, and 19 was morphologically identifiable. The notauli of species in this cluster are distinctly grooved, and there is striate sculpture on the mesoscutal lamella and medial mesoscutellum. The Synopeas specimen from Myanmar, which is included in this clade in some analyses, also has grooved notauli and striate mesoscutellar sculpture . Therefore, the grooved notauli group may represent a distinct species cluster within Synopeas . Further investigation is needed to understand the evolution of notaulus morphology with respect to Dolichotrypes and Sactog aster species, at least some of which have grooved notauli . Similar notauli are also found in Inostemma , an early diverging group , which could mean that grooved notauli represent the plesiomorphic state . Masner and Huggert (1989) applied the same polarity to this character for morphological systematics at the genus and subfamily level.

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112 Other small clusters b ear more subtle morphological similarities. Synopeas spp. 7 and 22 are characterized by a distinctly ovoid head in frontal view and a unidentate mandible . All other Synopeas species in the analysis have a bidentate mandible. Synopeas spp. 6 and 8 both have the lateral pronotum with microsculpture throughout , whereas the other species in the analysis have the lateral pronotum only partially sculptured . Synopeas spp. 1, 1b, and 13 have more extensive sculpturing on the metasoma than do most other species treated here, although this character is absent in the specimen from the UK, which belongs to the same cluster , and present in Syno peas sp. 6 , which is not closely related. In general, each species was reared from a single genus of host plant. The known distributions of the gall host plants are restricted to Papua New Guinea or to Australasia (Table 2 4), suggesting that the distrib ution of the gall midges and their parasitoids are similarly limited . The exception is Synopeas sp. 7, which was reared from two different plants in different genera, families, and orders. This could mean that Synopeas sp. 7 is a generalist parasitoid. If it is a specialist parasitoid, it is possible that it specializes on a generalist herbivore, although most species of gall forming Cecidomyiidae are monophagous (Dorchin et al. , 2019; Miller and Raman, 2019). More likely, i f Synopeas sp. 7 is a specialist, it attacks a predatory or inquiline midge that can be found on different plants. Synopeas sp. 7 emerged from at least five different gall morpho types identified by Philip Butterill, supporting the idea that it does not spe cialize on a gall former. The barcode data and morphology of Synopeas sp. 7 should be compared to that of Synopeas from other biogeographic regions before a formal description is published.

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113 The Synopeas species of New Guinea are not monophyletic , a result that lends itself to multiple interpretations. It is possible that there have been multiple introductions of Synopeas to the island. Alternatively, it could be that the region is a center of adaptive radiation for the genus Synopeas . The latter interpretat ion is in agreement with Masner and Huggert (1989), who suggested a Gondwanan origin for the Platygastrinae , at least in part . Drawing conclusions about deep evolutionary history will require more powerful tools than C OI and light microscopy . Such studies often integrate genomics, geometric morphometrics, 3D imaging, and other advanced techniques , as in Liu et al. (2020) . Although limited in its evolutionary relevance , the DNA barcoding component of the project has potential applications for biologi cal control and quarantine. Establishing a barcoding protocol and initiation of a database of Synopeas sequences will allow future researchers to better evaluate, monitor, and intercept adventive parasitoid species. With molecular methods, it will be possi ble to detect the presence of immature Synopeas within a cecidomyiid larva or gall, although barcoding alone cannot determine if the parasitoid is viable (Gariepy et al. , 2007). Barcoding cannot replace experimental host range testing in biological control programs because it does not measure developmental parameters . However , molecular data ca n enhance experimental , exploratory , and observational studies of parasitoid host relationships in agricultural and natural ecosystems. To achieve the full potential of such research will require many more barcode sequences from properly identified Synopeas species.

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114 CHAPTER 4 CONCLUSIONS Study Significance The results of my research emphasize the importance of certain tools to the integrative taxonomy of microhymenoptera. Despite the availability of more advanced molecular techniques, COI barcoding remains relevant and useful. Although the COI gene offers a relatively small amount of data and cannot provide a complete phylogenetic picture, its accessibility and convenience continue to make it an excellent standard barcode region for parasitic wasps. Morphology is essential to taxonomy, particularly when name bearing specimens have no associated molecular data, but also for understanding how organisms relate to one another and to their environments. Microphotography is an incredibly powerful tool for morphological analysis . High quality images can allow resear chers around the world to examine holotypes, identify specimens, and discuss characters. Beautiful images can also inspire interest and support f or entomology from the public . In the modern world, with its travel restrictions, socio economic inequalities, and near endless uncatalogued biodiversity, improving access to collections through digitized specimens is more important than ever before . Finally, the availability of reared material was crucial to the success of my project. Ecological data helped to def ine the likely geographic distribution of the described species and provided insight into parasitoid host relationships. Although rearing galls and larvae is challenging, reared parasitoid specimen s are more

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115 informative than those caught in traps or nets. Their value to integrative taxonomy cannot be overstated. Future Directions Future efforts may apply more advanced tools to Synopeas . Analysis of nuclear genes , ultra conserved elements , or genomics will provide more molecular data and could form the basis of more robust phylogenies. The results of such studies may closely resemble the story told by mitochondrial COI, supporting the clades and species groups suggested by my thesis research . However, different genes can evolve in different ways, and it is entirely possible that future molecular work will reveal a n alternative evolutionary history for Synopeas . My research only accounted for morphological characters visible with light microsco py. More morphological data may be found with scanning electron microscopy, computed tomography (CT) scanning, geometric morphometrics, and wing interference patterns. Even with light microscopy alone, there certainly remain characters to be found in Synop eas , especially on the legs, which I mostly ignored, and in the chaetotaxy, which shows obvious differences but has yet to be formalized. Genitalic dissection may also reveal valuable characters for species delimitation . The integrative taxonomy of the Syn opeas of New Guinea provides a foundation for revisionary work in other geographical regions. A full revision of Synopeas will have to begin in Europe, starting with examination of the oldest types from Förster, Walker, Westwood, and their contemporaries. The Palearctic region has a long history of species descriptions by isolated workers and is thus likely to include many synonyms. The Neotropical re gio n is a relatively clean slate. South Asia is a high priority region, as

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116 a revision is necessary to develo p identification tools for mango midge parasitoids. Finally, Nearctic Synopeas are ripe for revision due to good holotype access and the need to prepare for the gall midge problems of the future. We have only just begun to scratch the surface of the incre dible biodiversity of microhymenoptera. Parasitoid wasps are one of the most speciose groups in nature. Forbes et al. (2018) calculated that if two parasitoid wasp species occur for every five species of insect, there must be at least 883,000 species of Hy menoptera. Parasitoids of Cecidomyiidae are likely to be especially speciose, given that gall midges also have an extremely high biodiversity estimate, well over one million species (Hebert et al. , 2016). Although advancing human knowledge of the vast and mysterious Apocrita sometimes feels like emptying the ocean with a bucket, the need to understand our ecological and agricultural systems can help to prioritize parasitoid taxa for scientific attention. The diagnostic framework for Synopeas presented here is a small but significant step forward in the mission to identify, catalog, and monitor these beautiful and important little animals.

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117 LIST OF REFERENCES Abram PK, Haye T, Mason PG, Cappuccino N, Boivin G, Kuhlmann U ( 2012a ) Biology of Synopeas myles , a parasitoid of the swede midge, Contarinia nasturtii , in Europe. BioControl 57 : 789 800. Abram PK, Haye T, Mason PG, Cappuccino N, Boivin G, Kuhlmann U ( 2012b ) Identity, distribution, and seasonal phenology of parasitoids of the swede mid ge, Contarinia nasturtii (Kieffer) (Diptera: Cecidomyiidae) in Europe. Biological Control 62 : 197 205. Ashmead W H ( 1893 ) A m onograph of the North American Proctotrypidae. Bulletin of the United States National Museum 45 : 1 472 . Austin A D ( 1984 ) New speci es of Platygastridae (Hymenoptera) from India which parasitise pests of mango, particularly Procontarinia spp. (Diptera: Cecidomyiidae). Bulletin of Entomological Research 74 : 549 557. Austin AD, Johnson NF, Downton M ( 2005 ) Systematics, evolution, and bi ology of scelionid and platygastrid wasps. Annual Review of Entomology 50 : 553 5 82. Buhl PN (1997) On some new or little known species of Platygastrinae (Hymenoptera, Platygastridae). Entomofauna Zeitschrift für Entomologie 18: 429 468. Buhl PN (2004) New Australasian Platygastrinae (Hymenoptera: Platygastridae). Folia Entomologica Hungarica 65: 85 106. Buhl PN ( 2009 ) Taxonomical studies on Oriental Platygastridae (Hymenoptera: Platygastroidea). Journal of Asia Pacific Entomology 12 : 123 132. B uhl PN (2013) New or little known Platygastrinae (Hymenoptera: Platygastridae) from New Guinea. Monthly Magazine 149: 3 18 . Butterill PT, Novotny V ( 2015 ) Gall forming insects in a lowland tropical rainforest: Low species diversity in an extremely specialised guild. Ecological Entomology 40 : 409 419. Clausen CP ( 1978 ) Cecidomyiidae (Itonididae). In: Clausen CP (Ed) Introduced Parasites and Predators of Arthropod Pests and Weeds: A World Review. United States Department of Agriculture, Agricultural Research Service, Agriculture Handbook 4 80 : 316 318. Crawford JC, Bradley JC ( 1911 ) A new Pelecinus like genus and spec ies of Platygasteridae. Proceedings of the Entomological Society of Washington 13 : 124 125.

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125 BIOGRAPHICAL SKETCH Jessica Awad was born in Dayton, Ohio. As a teenager, s he moved to southern Florida, where she enrolled in Broward College . After earning an Associate of Arts degree , she transferred to the University of Florida Fort Lauderdale Research and Education Cente r. She supported herself with technician work while finishing her undergraduate degree in e ntomology and n ematology . After graduation, Jessica continued working as a professional laboratory technician. Her technician experience include s projects on ornamental horticulture, subterranean termites, plant parasitic nematodes, fire ant biological control, cactus moth biological control, museum specim en preparation , and fruit fly larval morphology . In 2016, Jessica positive first experience with parasitoid wasp identification. In 2017, she began her graduate studies as a non thesis student. Aft er a 2018 internship working with Synopeas , she decided she was n o t finished with the genus and needed to write a thesis on it. She is still not finished with Synopeas .