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The Biology of the Evolution of Viviparity in Asterinid Sea Stars

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The Biology of the Evolution of Viviparity in Asterinid Sea Stars THE BIOLOGY OF THE EVOLUTION OF VIVIPARITY IN ASTERINID SEA STARS Mohammad Sadequr Rahman Khan A thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy in the School of Life and Environmental Sciences, Faculty of Science, at The University of Sydney 2020 Dedicated to my dear parents and beloved wife DECLARATION I hereby declare that this work is my own, except where otherwise acknowledged. I also declare that this thesis has not been submitted in any form for another degree or diploma at any university or other institution. I consent to this thesis being made available for photocopying and loan under the appropriate Australian copyright laws. Mohammad Sadequr Rahman Khan Date: 10th February 2020 i AUTHORSHIP ATTRIBUTION Chapter 2 of this thesis is published: Khan, M.S.R., Whittington, C.M., Thompson, M.B., & Byrne, M. (2019) Intragonadal incubation of progeny in three viviparous asterinid sea stars that differ in offspring provisioning, lecithotrophy vs matrotrophy. Marine Biology, 166(6), 81. I analysed data and wrote the draft of the manuscript. MB Thompson and M Byrne assisted in sample collection. All co-authors contributed to manuscript preparation, revision and approved the final version of the manuscript. Chapter 3 of this thesis is partially published at Zoosymposia (2019), 15, 71–82 (see Appendix). I designed the study, collected and analysed data and wrote the draft of the manuscript. All co-authors assisted in research design, manuscript preparation and revision and approved the final version. Chapter 4 I did microscopy, analysed images, and wrote the manuscript. M Byrne prepared the histological slides. All co-authors assisted in manuscript preparation and revision. Chapter 5 of this thesis is partly published: Khan, M.S.R., Whittington, C.M., Thompson, M.B. & Byrne, M. (2019) Arrangement and size variation of intra-gonadal offspring in a viviparous asterinid sea star. Zoosymposia, 15, 71–82 (see Appendix). I did microscopy, analysed data and wrote the draft of the manuscript. All co-authors assisted in research design, manuscript preparation and revision of the manuscript and approved the final version for submission. ii As supervisor for the candidature upon which this thesis is based, we confirm that the authorship attribution statements above are correct. Prof. Michael B. Thompson Date: 10th February 2020 Prof. Maria Byrne Date: 10th February 2020 Dr. Camilla Whittington Date: 10th February 2020 iii ABSTRACT Viviparous asterinids exhibit great diversity in reproductive and offspring provisioning strategies, which raises fascinating life-history questions. This thesis investigates the biology of parent-offspring size variation, offspring release, nutrient provisioning and morphological adaptions in three viviparous asterinid sea stars, Cryptasterina hystera, Parvulastra vivipara and P. parvivipara. These species have contrasting lecithotrophic and matrotrophic provisioning of developing offspring in the gonads. In C. hystera (lecithotrophic), the juveniles (655 µm diameter) develop from large eggs (440 µm diameter). In P. vivipara and P. parvivipara, juveniles vary greatly in diameter (500–5000 µm) and develop from small eggs (84–150 µm diameter) through sibling cannibalism (matrotrophy). In these species, larger parents had greater reproductive output and produced more, but not larger, offspring. The species with matrotrophic offspring provisioning had a higher reproductive output than the lecithotrophic species. Parvulastra parvivipara released juveniles in 1–5 cohorts and exhibits continuous reproduction. Cryptasterina hystera retained a few large offspring in the gonad after 30 days of synchronous release. The degree of parental investment measured as matrotrophy index (the ratio of juvenile to egg dry mass) ranged from 597–55082 (P. parvivipara) and 1.7–6.2 (C. hystera), indicating a continuum in offspring provisioning. Potential specializations for viviparity and provisioning of nutrients for offspring were investigated using confocal microscopy and histology. The early larvae were closely associated with the inner gonad wall, supported by thin processes from somatic cells. The arrangement of P. parvivipara progeny in the gonads was observed three-dimensionally using micro-computed tomography. The juveniles were orally opposite to each other, presumably as a defensive strategy to protect themselves from being eaten. Confocal microscopy revealed 2–6 developmental stages in each gonad. The size variation of offspring intensifies when siblings start cannibalism post-metamorphosis. iv ACKNOWLEDGEMENTS It would have not been possible for me to write this thesis or publish my articles without the support of my talented supervisory team: Prof. Mike Thompson, Prof. Maria Byrne, and Dr. Camilla Whittington. My heartiest gratitude to Mike for choosing me as a PhD student and guiding me throughout this tough journey. Thanks, Mike, for your constructive criticism, inspiration, suggestions and numerous revisions of my writing. You were a great mentor for me. I have met very few people as smart as you are. Your financial supports were really helpful, as they allowed me to introduce myself to renowned researchers in national and international conferences. I wish you sound health. I am proud and honoured to have Maria Byrne as one of my supervisors. Her suggestions and active support in research design, data collection and innumerable revisions to prepare my manuscripts made it possible to submit this thesis on time. Maria, someone verily said that you are the grandma in the field of echinoderm research. Your expertise, supervision, data interpretation, and article writing pattern overwhelmed me and taught me a lot. There are very few supervisors who care for their students as much as you do. I wish you sound health and a long life. Camilla Whittington, my special thanks for your inspiration, support and guidance. You continuously pushed me towards achieving my goal. I am motivated by your enthusiasm and devotion for research. You were very kind to consider my food restrictions in every lab party. I am sure you will reach the peak of your expectations by dint of your quality, devotion and expertise. I express my gratitude to Liz McTaggart, Senior Natural Resources Officer, Department of Environment, Water and Natural Resources, South Australia, and to Dr. William Figueira and Mr. Geoff Prestedge for their assistance in sample collection. I would also like to thank Dr. Christopher Friesen, Dr. Mathew Crowther, and Dr. James Van Dyke for their assistance in data analyses. Thanks, also, to the Ministry of Agriculture, Fisheries and Forestry, Government of South Australia for the Ministerial Exemption to collect samples. I also acknowledge the Advance Microscopy Facility at the Bosch Institute and the Australian Centre for Microscopy and Microanalysis at the University of Sydney for allowing v me to use their equipment, and for the assistance provided by the staff of these facilities. I want to thank Prof. Chris Murphy, Matthew Foley, Sadaf Kalam, and Nasir Uddin for their assistance with lab techniques and research ideas. Obviously, the three and half years long journey would not be possible without continuous support, caring and feedback from my fellow lab members, especially Jacqueline Herbert, Jessica Dudley, Charles Foster, Claudia Santori, Alice Buddle, Zoe Skalkos, Melanie Laird, Monty Oldroyd, Josh Kemsley, Henrique Braz, and Oliver Griffith. I also thank Claudia Santori for taking care of the sea stars in my absence, and discussing a lot of issues. I will not forget the help that I received from Dr. Paula Cisternas, Januar Harianto, Dione Deaker and Hamish Campbell. I would like to thank HDRAC, the University Sydney for providing me with a fully funded University Sydney International Scholarship, without which my desire to carry out a PhD would have not been possible. Also thanks for awarding me PRSS grants for participating in conferences. I could not have felt more welcome in Sydney. Some people who did not know me before were really helpful and kindly opened their doors and let me stay in their safe and comfortable home. Of them, I want to mention Abdul Muktadir and Umme Salma. Also thanks to people that made my life abroad enjoyable. Last but not least, I would not be here if my parents had not supported me from my childhood. They are the best parents I could have ever asked for. My beloved wife was always beside me during most of my PhD and helped me a lot mentally by encouraging and supporting me in stressful situations. Every day, she cooked tasty foods even though she was pregnant and doing her own PhD works. Thanks Mansura for choosing me as your husband and staying with me in my woes and happiness. vi TABLE OF CONTENTS DECLARATION................................................................................................................................... I AUTHORSHIP ATTRIBUTION ....................................................................................................... II ABSTRACT .........................................................................................................................................IV ACKNOWLEDGEMENTS ................................................................................................................ V TABLE OF CONTENTS ................................................................................................................. VII LIST OF FIGURES ............................................................................................................................
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