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The Biology of the Cosmopolitan Fish Parasite Neobenedenia Girellae

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The Biology of the Cosmopolitan Fish Parasite Neobenedenia Girellae ResearchOnline@JCU This file is part of the following reference: Brazenor, Alexander Karlis (2017) The biology of the cosmopolitan fish parasite Neobenedenia girellae. PhD thesis, James Cook University. Access to this file is available from: http://dx.doi.org/10.4225/28/5a93846f8d373 The author has certified to JCU that they have made a reasonable effort to gain permission and acknowledge the owner of any third party copyright material included in this document. If you believe that this is not the case, please contact [email protected] The biology of the cosmopolitan fish parasite Neobenedenia girellae Thesis submitted by Alexander Karlis Brazenor BSc (Hons.) In fulfilment of the requirements for the Doctorate of Philosophy (Science) in the College of Marine and Environmental Sciences James Cook University Statement on the Contribution of Others Financial support for this study was provided by the College of Marine and Environmental Sciences, James Cook University (JCU). An Australian Postgraduate Award stipend provided support of living costs between 2012 and 2015. I was awarded a James Cook University Graduate Research School scholarship in 2014 to assist in paying for lab consumables and materials crucial to completing my experiments. The National Climate Change Adaptation Research Facility also awarded me with a grant that provided me with materials and lab consumables for conducting experiments. Travel grants to conferences were provided by the Australian Society for Parasitology, the Fisheries Research Development Corporation, and Fisheries Society of the British Isles. Most significantly, the Australian Society for Parasitology awarded me a travel grant in 2012 (covering flights and accommodation costs) to travel to Adelaide to begin my molecular analysis of parasite samples with Terry Bertozzi. Guidance in the analyses used in my thesis was provided by Kate Hutson, Richard Saunders, Nicholas Paul, Ross Alford, Martin Sheaves, and Rhondda Jones. Leslie Chisholm and Robert Adlard of the South Australia Museum and Queensland Museum respectively assisted in accessioning samples from Chapters 2, 3, and 4. Chapter Co-author Contribution Chapter 2 Terry Bertozzi Terry Bertozzi provided assistance in genetic analyses and editorial support. Terry Miller Terry Miller provided assistance in genetic analyses and editorial support. Ian Whittington Ian Whittington was instrumental in the development of ideas, the i provision of samples, and taxonomic identification of specimens. Kate Hutson Kate Hutson provided financial and editorial support and assisted in the development of ideas. Chapter 3 Richard Saunders Richard Saunders assisted in developing the aims of this paper, in the statistical analysis of the data, and in editing. Terry Miller Terry Miller assisted in the genetic analysis of samples included in this paper. Kate Hutson Kate Hutson assisted in financial support, editorial assistance, and the development of ideas. Chapter 4 Kate Hutson Kate Hutson provided assistance in finances, experiments, statistical analyses, and editing. Chapter 5 David Francis David Francis assisted in the biochemical analysis of samples in this study and provided editorial support. Kate Hutson Kate provided assistance in the development of ideas and editing. Guy Carton Guy Carton assisted in the development of ideas and the editing of this paper. Chapter 6 David Francis David Francis assisted in the biochemical analysis of samples in this study and provided editorial support. Guy Carton Guy Carton assisted in the development of ideas in this paper. Kate Hutson Kate provided assistance in the development of ideas, financial support, and editing. Title page image: Adult Neobenedenia girellae. Photograph taken by A. Brazenor. Image dimensions = 250 µm × 100 µm. ii Acknowledgements At the time of submitting this thesis, I will have spent almost six years in the Marine Parasitology Laboratory for my Honours and PhD research at James Cook University. During this time, I have had the great pleasure of learning from wonderful people. Every single co-author I worked with offered something new that I could learn from and I gained an immense amount of knowledge from researchers from incredibly diverse backgrounds. I am eternally grateful to Terry Bertozzi, Terry Miller, Richard Saunders, Ian Whittington, and David Francis who all made this thesis possible. Thank you to Ben Lawes, Simon Wever, and Andrew Thompson of the Marine and Aquaculture Research Facility. Their help sourcing laboratory space, equipment use, and general systems management has been instrumental in conducting a large number of the experiments in this thesis. Of course, one of the most important relationships a PhD candidate has is the one with their supervisor(s). I was incredibly fortunate that I had two supervisors who offered such different perspectives to situations. There were many situations in which this proved to be very helpful in developing solutions to problems or for motivating me when I needed it most. Firstly, my gratitude to Guy Carton, my co-supervisor, for providing advice and instruction throughout the thesis. His ability to clarify directions in which to progress and to facilitate helpful connections was invaluable to the completion of this PhD and I will always be grateful for his contribution to this thesis. iii Throughout my time in the Marine Parasitology Laboratory, my supervisor Kate has always been attentive and enthusiastic about my research. She has carefully instructed and directed my development as a researcher but, more importantly, also as a person. She is the most influential role model in my professional life and I owe all of the invaluable lessons and knowledge I have accrued in these past years to her skilful and dedicated guidance. I could not imagine having a better supervisor. It has been a pleasure to be able to work with her in her laboratory and to develop a friendship with her that is immensely important to me. Thank you, Kate. For so much. I owe so much to all of my friends who have supported me throughout this thesis. To Scott, Nina, Michael, Darcy, Angus, and Jack you made completing this journey a lot easier than it would have been otherwise. Thank you. To my colleagues in the Marine Parasitology Laboratory, you have all done a lot to support me throughout this endeavour and I am eternally indebted to you all for that. To Alejandro and Thane in particular, throughout the PhD we’ve weathered good times and bad, often with the judicious application of scotch. I can truly call both of you my brothers and your support has meant so much to me. Thanks, guys. It goes without saying that the successful completion of this thesis could not have occurred without the help of countless people including other researchers, administrative staff, industry contacts, and colleagues. My sincere gratitude to James Cook University for providing me with a foundation on which to build my skills and for allowing me to develop my skills as a marine parasitologist. iv My sincere gratitude to the Australian Society for Parasitology for providing me with funds to travel to South Australia, the Fisheries Society of British Ichthyology for awarding me a travel grant to be able to present my research in Spain, and to the Fisheries Research and Development Corporation (FRDC) for allowing me to present at their conferences in Cairns. Presenting my research to my peers was always nerve wracking but it was also one of the most rewarding experiences as a scientist. The organisations above made this a reality and I am so grateful for their help. My family deserve far more credit for the completion of thesis than I am able to convey. They have been incredibly supportive and encouraging throughout my education and are the reason that I am able to be doing what I love. Finally, to my loving fiancée, Madeleine. You are the most supportive partner I can possibly imagine having. Thank you for the encouragement, the support, and the patience you have had for me throughout this thesis. This thesis is dedicated to my family, my friends, and to those who took the time in their lives to support me throughout this journey. v Abstract Monogeneans are a fascinating parasite group to study. They can be harmful pathogens of finfish and display interesting relationships with their hosts making research on these species engaging from both ecological and economic perspectives. Neobenedenia (Family Capsalidae) Yamaguti, 1963, is a notable monogenean genus and is notorious for the large number of potential host species that the constituent parasite taxa are able to infect. The distinct lack of research on Neobenedenia biology and the diversity of species present in Australian waters presented me with an opportunity to conduct a series of research studies designed to improve understanding of this parasite genus. The aim of this thesis was to provide an in-depth investigation of Neobenedenia phylogenetics and detailed biology (including the growth, morphology, reproduction, and biochemistry) of the species in Australia. The first data chapter (Chapter 2) determined the phylogenetic relationships between 33 Neobenedenia isolates by amplifying three genes; two nuclear (H3 and 28S rDNA) and one mitochondrial (cytochrome b). Isolates were collected from a total of 23 host species and nine countries in both hemispheres and included 16 isolates from 12 host fishes in Australia. Representative samples for Neobenedenia melleni MacCallum, 1927, Neobenedenia pacifica Bravo-Hollis, 1971, and Neobenedenia longiprostata Bravo-Hollis, 1971, formed discrete clades and collectively accounted for seven of the 33 samples used in this study. The remaining 26 isolates formed a single clade genetically distinct from all representative specimens. Morphological observation of these 26 isolates confirmed that they were morphologically indistinguishable from N. melleni despite displaying clear genetic differences in the phylogenetic trees. This confirmed the suggestion by the scientific community that Neobenedenia girellae vi Hargis, 1955, a species that was synonymised with N. melleni in 1996, should be reinstated as its own taxon, a recommendation that is encouraged by the authors.
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