Elucidating the diversity of mid-eastern Queensland Pinwheel Snails (Eupulmonata: Charopidae) using aspects of shell morphology Author Holcroft, Lorelle Published 2018-04 Thesis Type Thesis (Masters) School School of Environment and Sc DOI https://doi.org/10.25904/1912/1217 Copyright Statement The author owns the copyright in this thesis, unless stated otherwise. Downloaded from http://hdl.handle.net/10072/380059 Griffith Research Online https://research-repository.griffith.edu.au Elucidating the diversity of mid-eastern Queensland Pinwheel Snails (Eupulmonata: Charopidae) using aspects of shell morphology. Lorelle Holcroft School of Environment and Science Griffith University Submitted in fulfilment of the requirements of the degree of Master of Philosophy April, 2018 DECLARATION This work has not previously been submitted for a degree or diploma in any university. To the best of my knowledge and belief, the thesis contains no material previously published or written by another person except where due reference is made in the thesis itself. Lorelle Holcroft Elucidating the diversity of mid-eastern Queensland Pinwheel Snails (Eupulmonata: Charopidae) using aspects of shell morphology. Lorelle Holcroft s5063935 1 | P a g e ACKNOWLEDGMENTS The research presented in this dissertation could not have been done without the help of many colleagues and friends who supported me in many ways during the last two years. Only the names of some of them can be listed here. Firstly, Griffith University and my supervisors, Professor Jane Hughes and Dr Chris Burwell for their unstinting support, feedback and encouragement. Thank you, Jane for skillfully ensuring the right mix of ‘supervision’ and ‘independent latitude’ in research. I have learnt much these past two years, and that could not have happened without your guidance. I have benefited greatly from the valuable feedback and suggestions of my other supervisor, Dr Chris Burwell. Thank you Chris, for the comprehensive critique and discussions. This was enormously informative and constructive. Thanks also to Kathryn Real from Griffith University for the DNA extraction and subsequent support and especially to PhD student, Jeremy Wilson, for help and guidance with the cladistic analyses of the molecular data. Sincere gratitude is expressed to two eminent biologists, Dr Christine Lambkin from the Queensland Museum who spent time teaching the process for running a PAUP analysis and who supervised the author’s first efforts, and Dr Frank Köhler from the Australian Museum for his support for the DNA analysis. Over the past two years, I have had the great pleasure of interacting with an incredible group of people including PhD students and professors from the Griffith School of Environment and Science - the ‘Geek’ group. I do hope you enjoyed your treats. In particular, I would like to thank Jeremy Wilson, Dr Dan Schmidt and Kathryn Real for taking an interest in my research and offering valuable advice. To the staff at Queensland Museum who have been so encouraging and supportive whenever I have had questions, Dr Christine Lambkin, Darryl Potter, Dr John Healy, Geoff Thompson, photographer extraordinaire, all the Level 6 scientists and in particular, the vertebrate crowd who gave me space in their lab - thank you. The copyright of all photographs of shells in Chapters 2 to 5 belongs to the Queensland Museum unless otherwise stated. The Australian Rivers Institute is recognised for many educational opportunities and ongoing support offered. Petney Dickson, a big thank you for cheerfully handling all administrative niggles, and the IT staff at Queensland Museum for their technical support and wizardry. Finally, I would like to thank Dr John Stanisic who has provided immeasurable help and support over the course of this work, for continual encouragement, for brainstorming ideas and theories for many hours and for teaching me much about these tiny snails as well as providing me many forums in which to learn. His advice and insights have been extremely helpful in shaping the research and I would like to dedicate this thesis to John. To many family members and friends have assisted throughout the production of this thesis in a number of ways, my thanks go to them. Elucidating the diversity of mid-eastern Queensland Pinwheel Snails (Eupulmonata: Charopidae) using aspects of shell morphology. Lorelle Holcroft s5063935 2 | P a g e ABSTRACT Species of Charopidae from mid-eastern Queensland in the collections of the Queensland and Australian Museums were reviewed primarily using aspects of shell morphology. Protoconch sculpture played a key role in this revision and 261 protoconchs were examined by scanning electron microscopy in order to develop a basic terminology that could be used in descriptions of genera and species. Eleven baseline patterns, some with subtle variations, were established. Eight of these are shown to be present in mid- eastern Queensland Charopidae. Studies of protoconch and teleoconch sculpture combined with shell coiling patterns showed that a number of genera and species, many newly documented, are represented. A cladistic analysis using a range of shell measurements was conducted to confirm these results. Additional verification, though limited, was sought through molecular analysis where animal tissue was available. Anatomical dissection of the male genitalia was performed on specimens of two species for which adequate preserved material was available. From an initial 13 known species, thirty-nine species of charopid are documented including a reallocation of the three ‘Gyrocochlea’ species currently described from the region resulting in three new genera, Whitcochlea gen. nov., Stanisicaropa gen. nov. and Radiolaropa gen. nov., and a new species, Radiolaropa eungella sp. nov. An additional four new Gyrocochlea-grade species are described in four new genera, Amfractaropa gen. nov., Comularopa gen. nov. Nodularopa gen. nov., and Xenoropa gen. nov. The genera Isolderopa Stanisic 2010 and Pereduropa Stanisic 2010 are expanded with the description of six new species. A new genus, Tristanoropa gen. nov. is diagnosed for the inclusion of three new species. Twelve additional species, some based on damaged specimens, are briefly described in order to complete the inventory. The majority of mid-eastern Queensland species appear to be endemic to the region. Biogeographical discussions focus on how the charopid fauna of the Central Mackay Coast Bioregion relates to similar faunas in bioregions to the north and south. Isolation of the region and its rainforests by dry corridors to the north and south that have developed as result of continental aridification since the Miocene may offer an explanation for the high endemicity of the mid-eastern Queensland charopid species. PAPERS INCLUDED IN THESIS Included in this thesis are papers in Chapters 3, 4 and 5 for which I am the sole author. Appropriate acknowledgements of those who contributed to the research but did not qualify as authors are included in each paper. The bibliographic details for these papers are: Chapter 3: Holcroft, L. 2018a. Protoconch sculpture as a taxonomic tool in Australian charopid systematics (Gastropoda: Eupulmonata: Charopidae), Molluscan Research, DOI:10.1080/13235818.2017.1409069. Molluscan Research copyright- permission for reprinting within the thesis. Elucidating the diversity of mid-eastern Queensland Pinwheel Snails (Eupulmonata: Charopidae) using aspects of shell morphology. Lorelle Holcroft s5063935 3 | P a g e Chapter 4: Holcroft, L. 2018b. A revision of Gyrocochlea-grade Charopidae from mid-eastern Queensland and a redescription and generic reassignment of Gyrocochlea iuloidea (Forbes, 1851), G. chambersae Stanisic 2010 and G. danieli Stanisic, 2010. (Eupulmonata: Charopidae). Memoirs of the Queensland Museum-Nature, DOI:10.17082/j.2204-1478.60.2018.2017-14. Queensland Museum- permission for reprinting within the thesis. Chapter 5: Holcroft, L. in prep. A revision of Charopidae with a finely cancellate protoconch sculpture from mid-eastern Queensland. (Eupulmonata: Charopidae). Memoirs of the Queensland Museum- Nature (in prep). TABLE OF CONTENTS DECLARATION ................................................................................................................................... 1 ACKNOWLEDGMENTS ....................................................................................................................... 2 ABSTRACT.......................................................................................................................................... 3 PAPERS INCLUDED IN THESIS ......................................................................................................... 3 TABLE OF CONTENTS ................................................................................................................... 4 DIAGRAMS AND ILLUSTRATIONS ................................................................................................. 9 LIST OF TABLES .......................................................................................................................... 11 CHAPTER 1: Introduction ................................................................................................................ 12 1.2 Aims of this study ......................................................................................................................... 13 1.3 Thesis overview ..........................................................................................................................