Systematics of Pseudoloxops Kirkaldy (Insecta: Heteroptera: Miridae)
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SYSTEMATICS OF PSEUDOLOXOPS KIRKALDY (INSECTA: HETEROPTERA: MIRIDAE) Ratnawati Hazali BSc. (Hons). (Animal Resource Science and Management) A thesis submitted in fulfillment of the requirements for the degree of Master of Science In Systematic Entomology School of Biological, Earth and Environmental Sciences Faculty of Science University of New South Wales 2013 PLEASETYPE THEUNIVERSIW OF NEWSOUTH WALES Thesis/DissertationSheet Surnameor Familyname: HAZALI Firstname: RATNAWATI Othername/s: Abbreviationfor degreeas givenin the Universitycalendar: M.Sc. School: Blologlcal, Earth and Environmental Sciences Faculty: Science Title: Systematlcs of Pseudoloxops Kirkaldy (lnsecta: Heteroptera: Miridae) Abdllct 450worda maxlmum:(PLEASE TYPE) The ganus PseuddarcF Kkkaldy (lnsocia:Heteroptera: Miridae: Orthotylinae: O.thotylinl) cunently comprises38 described sp€cissfrom ths Palaearctic,Oriental, Alrotroplcal and Australianregions, as well as islandsin the PacificBasin (Schuh1995, 20Og), and many new species await d€scription. Pseudoloxopslacks a comprehensive and modern systematlc treatnent, All prwious sfudia8 of Psatrtobxops ue rcgiond in scopg, and most authors have not evaluatedthe generic limits of Psetdoloxops acro88its dlstrlbutlonalrange. This proj€ctaddrsssed this taxonomicimpediment and the genericlimits of Psa.rdolo(ops.This syst€maticsudy was basedon an examinationot sp€cim€nsobtained lrom museumcollections, as well as n€w collections.A comparativ€ morphological study based on €xtsrnal and male genitalia characters has been done to identify characters ol dlagno8dcand phylogeneticimportenc€. A phylogeneticanalysis of morphologicaldataset ot 53 taxa and 78 charact€rsis preesnt€d,The r€sults sho$, that the genlA Psaudola<opsis r€cognisedas a monophyleticgroup based on synapomorphiesot the head. The Southwest Pacific group of 20 sp€cies was selected ior further systematic revision. A total ol 19 new sp€cies are d€scrlbsdlncludlng the one previouslydesctibed sp€cies, Fseudoloxops vitiercis Kirkaldy. Habltus color photographs, illustrations oJths mal€genitalia and distributionmaps ars provid€d. Declaratlonrelatlng to dlsposltlon thesis/dissertation I herebygrant to the Universityof NewSouth Wales or its agentsthe rightto archiveand to makeavailable my thesisor dissertationin wholeor in partin the Universitylibraries in all formsof media,now or hereafter known, subject to the provisionsof the CopyrightAct 1968.I retainall propertyrights, such as patentrights. I alsoretain the rightto use in futureworks (such as articlesor books)all or partof thisthesis or dissertation. I alsoauthorise University Microfilms to usethe 350word abstract of my thesisin Dissg lion Abstraaslnternational (this is applicableto doctoral thesesonly). t/ /og/roir .'.'.'.r.'...'....\r...'....'.. Date The Universityrecognises that there may be exceptionalcircumstances requiring restrictions on copyingor conditionson use. Requestsfor restrictionfor a periodof up to 2 yearsmust be madein writing. Requestsfor a longerperiod of restrictionmay be consideredin exceptional circumstancesand requirethe of the Deanof GraduateResearch. FOROFFICE USE ONLY Dateof completionof requirementsfor Award: THISSHEET IS TO BE GLUEDTO THE INSIDEFRONT COVER OF THE THESIS ORIGINALIWSTATEMENT 'l herebydeclare that this submissionis my own work and to the best of my knowledgeit containsno materialspreviously published or writtenby another person,or substantialproportions of materialwhich have been acceptedfor the award of any other degree or dlploma at UNSW or any other educational institution,except where due acknowledgementis made in the thesis. Any contributionmade to the research by others, with whom I have worked at UNSWor elsewhere,is explicitlyacknowledged in the thesis.I also declarethat the intellectualcontent of this thesis is the productof my own work, except to the extentthat assistancefrom others in the project'sdesign and conceptionor in style,presentation and linguisticexpression is acknowledged.' COPYRIGHTSTATEMENT 'l herebygrant the Universityof New SouthWales or its agentsthe right to archiveand to makeavailable my thesisor dissertationin wholeor part in the Universitylibraries in all formsof media,now or hereafter known, subject to the provisionsof the CopyrightAct 1968.I retainall proprietaryrights, such as patent rights.I also retainthe rightto use in futureworks (such as articlesor books)all or partof thisthesis or dissertation. I alsoauthorise University Microfilms to usethe 350 wordabstract of my thesisin DissertationAbsfract International (this is applicableto doctoralthesesonly). I have eitherused no substantialportions of copyrightmaterial in my thesisor I haveobtained permission to use copyrightmaterial; where permission has not beengranted I haveappliedlwill apply for a partialrestriction of the digitalcopy of mythesis or dissertation.' AUTHENTICIW STATEMENT 'l certiff that the Library deposit digital copy is a direct equivalentof the final officially approved version of my thesis. No emendation of content has occurred and if there are any minor variations in formatting, they are the result of the conversionto digitalformat.' Date */::frt? ACKNOWLEDGEMENTS First and foremost, I would like to thank Universiti Malaysia Sarawak (UNIMAS) and the Malaysian Ministry of Higher Education for granting me the scholarship for my graduate study at University of New South Wales. Deepest thanks to my supervisor, Professor Gerry Cassis, without whose support, guidance and faith, this thesis would not have been completed. I’m definitely proud to be one of his students and grateful to him for accepting me as his student and educating me. I thank Professor Mike Archer for introducing me to Gerry. Without him I would not be in Prof. Cassis’ lab today. There are many individuals who have been part of my project. Importantly, I would like to thank Celia Symonds, Hannah Finlay and Dr. Nik Tatarnic for their invaluable assistance from the beginning of this project. I appreciate all their time, expertise, guidance and friendship. Special thanks to my lab members particularly Fiona Powell, Michael Elias, Anna Namyatova, Marina Cheng and Alex Brown. They have been great friends, providing an enjoyable work environment and support throughout this whole process. I would like to acknowledge those from many institutions around the world who generously provided specimens for this study. I would also like to thank the Evolution & Ecology Research Centre (E&ERC), School of Biological, Earth and Environmental Sciences for the administrative and financial assistance given throughout the last semester of my study. i I give a special thank you to Professor Fatimah Hj. Abang for believing in me and for her continual support. Finally, I would like to acknowledge my parents, Mr. Hazali Hj. Aman and Mrs. Rosiah Sebli, and my siblings for their love and unwavering support. Special thanks go to Wahap Marni for his constant encouragement. Last but not least, sincere thanks are given to all the many people who I have not listed here, but who have helped me in one way or another. I am very fortunate to have such wonderful and loving people around me. ii ABSTRACT The genus Pseudoloxops Kirkaldy (Insecta: Heteroptera: Miridae: Orthotylinae: Orthotylini) currently comprises 38 described species from the Palaearctic, Oriental, Afrotropical and Australian regions, as well as islands in the Pacific Basin (Schuh 1995, 2008), and many new species await description. Pseudoloxops lacks a comprehensive and modern systematic treatment. All previous studies of Pseudoloxops are regional in scope, and most authors have not evaluated the generic limits of Pseudoloxops across its distributional range. This project addressed this taxonomic impediment and the generic limits of Pseudoloxops. This systematic study was based on an examination of specimens obtained from museum collections, as well as new collections. A comparative morphological study based on external and male genitalia characters has been done to identify characters of diagnostic and phylogenetic importance. A phylogenetic analysis of morphological dataset of 53 taxa and 78 characters is presented. The results show that the genus Pseudoloxops is recognised as a monophyletic group based on synapomorphies of the head. The Southwest Pacific group of 20 species was selected for further systematic revision. A total of 19 new species are described including the one previously described species, Pseudoloxops vitiensis Kirkaldy. Habitus color photographs, illustrations of the male genitalia and distribution maps are provided. iii TABLE OF CONTENTS ACKNOWLEDGEMENTS ……………………………………………………….. i ABSTRACT ………………………………………………………………………. iii LIST OF TABLES ………………………………………………………………… v LIST OF FIGURES ……………………………………………………………….. vi CHAPTER I. GENERAL INTRODUCTION …………………………………….. 1 CHAPTER II. PHYLOGENETIC ANALYSIS OF THE GENUS PSEUDOLOXOPS KIRKALDY (INSECTA: HETEROPTERA: MIRIDAE: ORTHOTYLINAE) ……………………………………………………………….. 7 Introduction …………………………………………………………………. 7 Materials and methods ………………………………………………………. 9 Character homology and terminology ………………………………………. 12 Phylogenetic results …………………………………………………………. 17 Discussion …………………………………………………………………… 28 References …………………………………………………………………... 31 Tables and Figures …………………………………………………………... 37 CHAPTER III. A SYSTEMATIC STUDY OF THE GENUS PSEUDOLOXOPS KIRKALDY IN THE SOUTHWEST PACIFIC ………..……………………….. 54 Introduction …………………………………………………………………. 54 Materials and methods ………………………………………………………. 56 Taxonomy ……………………………………………………………………