TAXONOMY AND PHYLOGENETICS OF THE GENUS ULONEMIA DRAKE AND POOR, 1937 (HETEROPTERA: TINGIDAE), WITH AN EMPHASIS ON THE POPULATION GENETICS OF A PESTIFEROUS SPECIES Ryan M. Shofner A thesis in fulfilment of the requirements for the degree of Doctor of Philosophy School of Biological, Earth and Environmental Sciences Faculty of Science August 2018 Thesis/Dissertation Sheet Surname/Family Name : Shofner Given Name/s : Ryan Michael Abbreviation for degree as give in the University calendar : PhD Faculty : Science School : Biological, Earth, and Environmental Sciences Taxonomy and phylogenetics of the genus Ulonemia Drake and Poor, 1937 Thesis Title : (Heteroptera: Tingidae), with an emphasis on the population genetics of a pestiferous species Abstract 350 words maximum: (PLEASE TYPE) Agricultural systems are increasingly faced with the emergence of novel pests. Within the last decade, the macadamia lace bug, Ulonemia decoris Drake, 1942, became established in macadamia orchards in northern New South Wales and southern Queensland. This study provides the first comprehensive overview of the genus and its status in Australia. This study had three broad goals: 1) to describe the relationship between Ulonemia and other tingids in Australia using genetics; 2) to redefine the genus Ulonemia using the results from part 1; 3) to examine the population genomics of U. decoris to determine the dispersal rate of individuals between populations. Analysis of four genes (two mitochondrial and two nuclear) by Maximum Likelihood and Bayesian estimation returned a paraphyletic Ulonemia with three strongly supported clades; a combined morphological and molecular analysis with parsimony failed to resolve any relationships. Two new genera and five new species, Cercotingis (C. croajingolong sp. nov., C. namadgi sp. nov., C. tasmaniensis sp. nov.) and Proteatingis (P. astibosetes sp. nov., P. howardi sp. nov.), are described to account for the observed molecular phylogenies. One species of Tingis and U. decoris were transferred into Cercotingis, and three species of Ulonemia were transferred to Proteatingis. The capability of lace bugs to disperse between orchards is critical in developing better control methods, because determining dispersal ability can provide valuable information on the geographic extent over which control must be coordinated. Single-nucleotide polymorphism (SNP) data were obtained for C. decoris for 204 individuals over 7 localities in northern NSW. Each sample locality had an excess of homozygous individuals and there was minimal genetic differentiation over geographic distance. This evidence points to selection against heterozygotes, recent extensive mixing, non-random mating, parthenogenesis, or cryptic species within C. decoris. The high dispersal and possible rapid reproduction due to parthenogenesis would make C. decoris difficult to manage. There is also a need to monitor for other emergent pest species, because there are multiple lace bug lineages on the Proteaceae, with three of these lineages containing verified pests. In summary, optimal management of the species will likely require region-wide coordination. Declaration relating to disposition of project thesis/dissertation I hereby grant to the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or in part in the University libraries in all forms of media, now or here after known, subject to the provisions of the Copyright Act 1968. I retain all property rights, such as patent rights. I also retain the right to use in future works (such as articles or books) all or part of this thesis or dissertation. I also authorise University Microfilms to use the 350 word abstract of my thesis in Dissertation Abstracts International (this is applicable to doctoral theses only). …………………………………………………………… ……………………………………..……………… ……….……………………...…….… Signature Witness Signature Date The University recognises that there may be exceptional circumstances requiring restrictions on copying or conditions on use. Requests for restriction for a period of up to 2 years must be made in writing. Requests for a longer period of restriction may be considered in exceptional circumstances and require the approval of the Dean of Graduate Research. FOR OFFICE USE ONLY Date of completion of requirements for Award: ORIGINALITY STATEMENT ‘I hereby declare that this submission is my own work and to the best of my knowledge it contains no materials previously published or written by another person, or substantial proportions of material which have been accepted for the award of any other degree or diploma at UNSW or any other educational institution, except where due acknowledgement is made in the thesis. Any contribution made to the research by others, with whom I have worked at UNSW or elsewhere, is explicitly acknowledged in the thesis. I also declare that the intellectual content of this thesis is the product of my own work, except to the extent that assistance from others in the project's design and conception or in style, presentation and linguistic expression is acknowledged.’ Signed …………………………………………….............. Date …………………………………………….............. COPYRIGHT STATEMENT ‘I hereby grant the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or part in the University libraries in all forms of media, now or here after known, subject to the provisions of the Copyright Act 1968. I retain all proprietary rights, such as patent rights. I also retain the right to use in future works (such as articles or books) all or part of this thesis or dissertation. I also authorise University Microfilms to use the 350 word abstract of my thesis in Dissertation Abstract International (this is applicable to doctoral theses only). I have either used no substantial portions of copyright material in my thesis or I have obtained permission to use copyright material; where permission has not been granted I have applied/will apply for a partial restriction of the digital copy of my thesis or dissertation.' Signed ……………………………………………........................... 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Publications can be used in their thesis in lieu of a Chapter if: • The student contributed greater than 50% of the content in the publication and is the “primary author”, ie. the student was responsible primarily for the planning, execution and preparation of the work for publication • The student has approval to include the publication in their thesis in lieu of a Chapter from their supervisor and Postgraduate Coordinator. • The publication is not subject to any obligations or contractual agreements with a third party that would constrain its inclusion in the thesis Please indicate whether this thesis contains published material or not. This thesis contains no publications, either published or submitted for publication Some of the work described in this thesis has been published and it has been documented in the relevant Chapters with acknowledgement This thesis has publications (either published or submitted for publication) incorporated into it in lieu of a chapter and the details are presented below CANDIDATE’S DECLARATION I declare that: • I have complied with the Thesis Examination Procedure • where I have used a publication in lieu of a Chapter, the listed publication(s) below meet(s) the requirements to be included in the thesis. Name Signature Date (dd/mm/yy) Postgraduate Coordinator’s Declaration (to be filled in where publications are used in lieu of Chapters) I declare that: • the information below is accurate • where listed publication(s) have been used in lieu of Chapter(s), their use complies with the Thesis Examination Procedure • the minimum requirements for the format of the thesis have been met. PGC’s Name PGC’s Signature Date (dd/mm/yy) i ACKNOWLEDGEMENTS This thesis was a long journey, possible only through the endless contributions of numerous wonderful people. I was fortunate to cover vast expanses of Australia in my quest for knowledge on lace bugs, and I have lost count of how many truly helpful and inspiring people I’ve met along the way. They have all contributed, in a small part at least, to this thesis, and all have earned my gratitude and thanks. First and foremost, I thank my supervisors, Gerry Cassis and Bill Sherwin, without whom this project would not have happened at all. They endured countless hours of questions as well as rebuttals when I thought I knew better. Without fail, they put be back on the correct track. I thank my committee Russell Bonduriansky, Shaun Laffin, and Terry Ord for the help and guidance they provided throughout my candidature. I thank Marina Cheng, without whom the project, and myself, would have fallen into a completely disorganised mess. From providing lab training, support, and logistics, or helping me in the field, to making sure I and everyone in