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Lineages of Hepialid Moths in New Zealand

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Lineages of Hepialid Moths in New Zealand Lincoln University Digital Thesis Copyright Statement The digital copy of this thesis is protected by the Copyright Act 1994 (New Zealand). This thesis may be consulted by you, provided you comply with the provisions of the Act and the following conditions of use: you will use the copy only for the purposes of research or private study you will recognise the author's right to be identified as the author of the thesis and due acknowledgement will be made to the author where appropriate you will obtain the author's permission before publishing any material from the thesis. THE- EVOLUTIONARY HISTORY OF THE 'OXYCANUS' LINEAGES OF HEPIALID MOTHS IN NEW ZEALAND A thesis submitted in partial fulfilment of the requirements of the Degree of Doctor of Philosophy at Lincoln University by Barbara Brown Lincoln University 1998 II Abstract of a thesis submitted in partial fulfilment of the requirements for the Degree of Ph.D. The evolutionary history of the 'Oxycanus' lineages of hepialid moths in New Zealand Barbara Brown This thesis attempted to reconstruct the evolutionary history of part of New Zealand's hepialid fauna. Phylogenetic analysis was used to test the monophyly of the informal 'Oxycanus' Cladoxycanus and 'Oxycanus' s. str. lineages. Data were morphological characters and molecular characters from the mtDNA COl & II and nrDNA ITS2 regions. Morphological and COl & II phylogenies indicated that the 'Oxycanus' Cladoxycanus and 'Oxycanus' s. str. lineages were monophyletic clades. Addition of Australian hepialid exemplars to the morphological data set indicated that Oxycanus and Jeana taxa were sister group to the monophyletic New Zealand 'Oxycanus' lineage s. str., while addition to the COl & II data set indicated that 'Oxycanus' Cladoxycanus was sister group to the 'Oxycanus' lineage s. str. The morphology and COl & II phylogenies were mostly congruent, with both recovering clades that corresponded to the informal Aenetus, Aoraia, 'Oxycanus' Cladoxycanus and 'Oxycanus's. str. lineages. The morphological and COl & II phylogenies were not congruent with respect to the placement of the Dumbletonius taxa. The morphological data set recovered Dumbletonius characterifer in a clade with D. unimaculatus, while the COl & II data set recovered D. characterifer as the basal taxon in the 'Oxycanus' lineage s. str. and D. unimaculatus in a clade with Wiseana taxa. Tracing the evolution of morphological characters that supported the monophyly of the genus Dumbletonius on to the independently derived COl & II phylogeny indicated that all characters were plesiomorphic. Spectral analysis gave higher support to the (D. unimaculatus, Wiseana) clade compared with the (D. characterifer, D. unimaculatus) clade. Morphological characters were mapped on to the COl & II phylogeny. Aenetus and Aoraia taxa were hypothesised to exhibit ancestral character states, male genitalic characters were less homoplasious compared with other data partitions and the morphological synapomorphies supporting the 'Oxycanus' lineages and the 'Oxycanus' s. str. lineage alone, were homologous. iii The combined morphology, COl & II and ITS2 data set did not support the monophyly of the 'Oxycanus' lineages. Dumbletonius unimaculatus, D. characterifer, Cladoxycanus, Heloxycanus and Dioxycanus taxa were recovered in a clade together. This topology was not recovered from any other data sets. Saturation of nucleotide sites in more divergent taxa and inconsistency in the parsimony method may have contributed to this result. The combined morphology and COl & II data set recovered a phylogeny congruent with that from the morphology alone. A combined morphology, COl & II, ITS2 and allozyme data set with Dumbletonius unimaculatus as outgroup, recovered the most resolved phylogeny for the genus Wiseana. The two clades recovered were: (C(Wiseana cervinata 'southern', Wiseana cervinata 'northern'), W. Jocosa), (W. copularis (southern', W. copularis 'northern'), (W. fuliginea, W. mimica)) and «W. signata 'southern', W. signata 'northern'), W. umbraculata). A diagnostic test was developed for Wiseana taxa based on the cleavage of the mtDNA COl & II gene regions with restriction enzymes. All Wiseana taxa, apart from W; fuliginea and W. mimica, were identified by unique restriction fragment length polymorphism patterns. Key words· hepialid, (Oxycanus' Cladoxycanus, (Oxycanus's. str., Aenetus, Aoraia, phylogeny, morphology, mtDNA COl & II, nrDNA ITS2, combined analysis, total evidence, taxonomic congruence, character mapping, diagnostic test, restriction fragment length polymorphism, restriction enzymes. iv ACKNOWLEDGEMENTS You probably won't believe me, but I have enjoyed every moment of this project. What a great chance to study a topic in depth and end up with more questions, get to know some real characters, visit scenic spots and develop new eccentricities. Not only has this been a productive time but it has also been creative. We now have the hepialid placemats©, the Wiseana dartboard©, the PhD snakes and ladders© and some wonderful new turns of phrase, e.g., lepidoptember (that month when the thesis was meant to be finished) and UUTrs and TUTrs (utterly useless and totally useless terminal taxa). Many people have contributed to the success of this project. In the Ecology and Entomology Group (formerly the Department of Entomology and Animal Ecology), thanks to Bruce Chapman and Eric Scott, as Heads of Department, for their encouragement and support. Thanks to my supervisors Rowan Emberson and Adrian Paterson for always being so willing to answer questions, discuss ideas and read drafts. Thanks for all the papers, books, moths, friendly chats, good advice, good parties, black jelly beans and pies. I really appreciate that Sue Worner, Sue Unsworth and Bruce Chapman took over my teaching duties while I had a year of study leave. Karen Armstrong and Charlotte Cameron were supportive in the lab and generous with their time, ideas, pipette tips, alcohol and film. The Lab Rats Lunch Club provided me with something other than ependorf tubes and lead pipes to chew on - thanks Charlotte, Helen, Tina, Dave and Anthony. Thanks to Bronwyn Hamilton and Diane O'Connor for helping me with those 'fiddly bits' of word processing. Special thanks to John Dugdale who instructed me in the finer points of hepia lid morphology and was generous with his time, books, drawings and great knowledge of the New Zealand lepidopteran fauna. John's work in Fauna ofNew Zealand 30, the oft­ quoted "Dugdale, 1994", was a valuable and inspiring resource for this study. Thanks to the organisations that provided funding for this study. I was privileged to be supported by the Lincoln University New Developments FU!ld, the Miss E.L. Hellaby Indigenous Grasslands Research Fund and the New Zealand Federation of University Women. v Thanks to all those who collected specimens for me. First prize for ingenuity goes to the person, who after running out of ethanol, sent me a specimen in gin because they didn't want to waste their whisky! Thanks to • Mike Bowie, Sue Britain, Jenny Brown, Muriel Brown, Libby Burgess, the Burridge's ofMasterton, Hilary Cooper, Janine Duckworth, John Dugdale, Rowan Emberson, Bruce Fraser, George Gibbs, Sonke Hardersen, Tina Lahmann, John Marris, Helen & Alastair McLees, Adrian Paterson, Brian Patrick, Peter Peckham, Katrin Schops, Geoff Spearpoint, Owen Spearpoint, Warren Thomas, Jackie Townsend, Ian Townsend and Graeme White. Ted Edwards and Margaret Williams from Australia provided specimens of Jeana timeata, Oxycanus diremptus and Trictena argentata. Thanks to Peter McQuillan and family, and Karyl Micheals in Hobart for their hospitality and logistical support during my field trip to Tasmania. Many people spent hours out in the field with me, huddled round the generator exhaust, trying to keep warm. Thanks to Mike Bayley, Tracey Bourner, Sven Brabyn, Lars Brabyn, Jenny Brown, John Dugdale, Alex Garnham, Dave Glenny, Lynette Hartley, Liz Herrick; Mark Pickering, Owen Spearpoint and Geoff Spearpoint. Thanks to all my family and friends who have supported me throughout this study. Geoff Spearpoint deserves a medal for his contribution. He was always calm when faced with fieldwork anxiety, not jealous when I was spending more time with the PCR machine and herculean in his efforts carrying the generator up on to the Craigieburn Range, in the fruitless search for Dioxycanus oreas. VI CONTENTS PAGE Title i Abstract ii Acknowledgements iv Table of Contents VI List of Tables Xl List of Figures XlI Chapter 1 Introduction 1 Species concepts 2 Characters 3 Tree building methods 4 Parsimony 4 Maximum likelihood 4 Accuracy of methods 5 Resolving conflict - within tree building methods 6 Resolving conflict - between tree building methods 6 Non-tree building methods of inferring phylogeny 6 Testing evolutionary hypotheses 7 The evolutionary history of the (Oxycanus' lineages of hepialid moths in New Zealand 7 Mms 8 Structure of the thesis 8 References 10 Chapter 2 Phylogeny of New Zealand hepialid moths (Lepidoptera: Hepialidae) inferred from a cladistic analysis of morphological data) Abstract 17 Introduction 18 Current classification of the New Zealand genera and species 19 Material and methods 22 Characters 22 Cladistic analysis 22 Terminal taxa and data matrix 22 Outgroups 23 viii Conclusions 83 Acknowledgements 83 References 84 Appendix 1 92 Appendix 2 94 Appendix 3 95 Appendix 4 101 Appendix 5 103 Chapter 4 Phylogeny of the New Zealand 'Oxycanus' lineages of hepialid moths (Lepidoptera: Hepialidae) inferred
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