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Biology and Behaviour of New Zealand Ladybirds

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Biology and Behaviour of New Zealand Ladybirds Biology and behaviour of New Zealand ladybirds What factors affect the successful establishment of introduced species? Our understanding of establishment processes is limited and concentrated on mean trait differences. However, recent research suggests that high levels of variation in functionally important traits facilitate establishment success. Ladybirds are generalist predators that serve as an excellent model system to study establishment success as they are widely introduced as biocontrol agents against agricultural pests. In my PhD, I examined the phenotypic (morphological, behavioural and physiological) variation and phylogenetic relationships between introduced and native ladybirds in New Zealand. Contrasting the differences between successfully established introduced species and native species in recipient ecosystems provides insights about the mechanisms that drive establishment success and a foundation for future research into introduction biology. Firstly, I developed a framework to study establishment processes using ladybirds as a model system. I identified fundamental gaps in research into unsuccessful species introductions, dispersal abilities and the quantification of both inter- and intraspecific variation in ladybirds. This provided the conceptual foundation for my subsequent research comparing inter- and intraspecific variation in the morphology, behaviour and physiology of New Zealand ladybirds. Morphologically, I found that introduced ladybirds had wider bodies than native ladybirds. Further, introduced species had greater intraspecific variation in size than native species. Introduced ladybirds with wide bodies and high intraspecific variation may have a larger prey range compared to native ladybirds, thereby increasing the probability of successful establishment. Behaviourally, I found differences between species in their general activity levels and responses to simulated predator attacks but there was no difference between species in their intraspecific variation. Physiologically, there was inter- and intraspecific variation in survival during desiccation resistance assays. Overall, body size appears to influence traits associated with establishment success. There was a strong positive correlation between body size and general activity i Biology and behaviour of New Zealand ladybirds levels and survival time during desiccation resistance assays in each species. I found that the widely dispersed introduced ladybird Halmus chalybeus had significantly higher levels of intraspecific variation in size and colour compared to other species of ladybirds. This suggests that high intraspecific variation may indeed increase establishment success and dispersal. Finally, the molecular phylogeny I constructed strongly supported the monophyly of Coccinellidae and subfamilies Microweiseinae and Coccinellinae and moderately supported the monophyly of tribe Epilachnini and paraphyletic tribes Coccinellini, Scymnini, Chilocorini and Coccidulini. However, there was no indication that the successful establishment of introduced species of ladybirds increases with relatedness to native species in New Zealand. This is the first study to compare phenotypic variation between introduced and native ladybirds in New Zealand and the first molecular phylogeny to include native species of New Zealand ladybirds. My thesis presents indirect support for the hypothesis that high levels of intraspecific variation increase establishment success in introduced species. I propose that assessing phenotypic variation in introduced species is key to understanding establishment processes and thereby improving the effectiveness of biocontrol and conservation programmes. ii Biology and behaviour of New Zealand ladybirds I would like to express my deepest appreciation to many wonderful people who made my PhD journey special and memorable. Firstly, I am extremely grateful for my amazing panel of supervisors. My principal supervisor Dr Anne Wignall has always been such an inspiration to me. From the day I first arrived in New Zealand she has always been there for me and guided me within and beyond the academic environment. I am immensely thankful for her help with my fieldwork as well. Dr Anne Wignall has always encouraged and motivated me to become a better scientist and researcher every day. I am extremely blessed to have been guided and mentored by Dr Anne Wignall and couldn’t have imagined having a better mentor for my PhD. I am also extremely thankful to all my co-supervisors, Dr Richard Leschen for his kind words of encouragement, motivation and for providing me with valuable feedback throughout my PhD. I greatly value your help with species identification and I enjoyed extremely the time I worked with you at Manaaki Whenua - Landcare Research. I would also like to thank Prof. James Dale for his guidance and continuous support over the past four years. I am extremely thankful for your guidance and insightful comments throughout my PhD. My sincere gratitude goes to Prof Dianne Brunton and Dr Luis Ortiz-Catedral for providing kind and encouraging feedback as my advisory committee at the confirmation seminar. A special thank you to Dr Aaron Harmer and Dr Daniel Thomas for their support during behavioural experiments to understand the use of the ‘PathtrackR’ library. I am also extremely thankful to Dr Libby Liggins for her guidance during phylogenetic data analysis. Attending the Molecular Ecology conference and Ira-Moana workshop was immensely helpful for me. I am extremely thankful to Dr Nikki Freed for her support during the desiccation resistance experiment to use the incubator in her lab. I am extremely grateful for the staff at the EcoGene lab at Manaaki Whenua; Talia Brav-Cubitt for guiding, training and assisting the sequencing process, Dr Robin Howitt and Duckchul Park for providing feedback and offering advice and guidance iii Biology and behaviour of New Zealand ladybirds when needed. Dr Thomas Buckley provided advice and support throughout the process. I also thank Grace Hall and Birgit Rhode for their assistance in the imaging and specimen access in the New Zealand Arthropod Collection (NZAC). I thank Manaaki Whenua – Landcare Research for allowing me to use specimens from the NZAC. My gratitude goes to all the funding sources including, the School of Natural and Computational Sciences - Massey University, Royal Society Te Apārangi and the Entomological Society of New Zealand for providing financial support for my PhD. I am extremely thankful for all my field buddies, Jin Choy, Nishadi Anuruddika, Jessica Patino Perez, Hoin An, Odette Howarth and Jessica Hiscox for their assistance in the field collecting ladybirds. I am also thankful for Hayden Pye and Charley Kho for their assistance in scoring images of ladybirds. I would like to thank Dr Asha Chaggan and Dr Gonzalo Avila from Plant and Food Research and Dr Darren Ward from Manaaki Whenua (Landcare Research), for meeting with me to educate me about the ongoing research of ladybirds in New Zealand. I would like to thank Alan Finn (Manaaki Whenua – Landcare Research) for sharing his expertise and experience with ladybirds and sharing his masters degree thesis with me for reference. I am also thankful for Dr Nicholas Martin (Manaaki Whenua – Landcare Research) for his insights into fieldwork and finding ladybirds at different stages of my PhD. My sincere appreciation to all the technical and administration staff including Annette Warbrook, Linh Mills, Cynthia Sim, Sue Di Leo, Mike Yep and Anil Malhotra for their assistance throughout my PhD. My sincere gratitude goes to all my past teachers and lecturers, especially to Dr Chandima Dangalle and Prof Thirimanna from University of Colombo, Sri Lanka for their words of encouragement. I am extremely grateful to all my wonderful PhD friends, Abigail Kuranchie, Akshya Ilangovan, Enzo Rodriguez-Reyes, Jessica Patino Perez, Mehrnaz Tavasoli, Michelle Roper, Simone Giovanardi and Wesley Webb for their constructive feedback, encouraging conversations and kind words of advice throughout my PhD. iv Biology and behaviour of New Zealand ladybirds I feel blessed to have all my friends and family in Sri Lanka, New Zealand and Australia for their love, support and encouragement. I am forever grateful to my mother, for your endless sacrifices, guidance and eternal love you gave me and wherever you are I feel your blessings every day, to my father, for always providing me with what I ask for and more, encouraging me and for teaching me how to accept challenges with a smile on my face and to my little brother for always being there for me. A huge thank you for my aunt Anoma and uncle Ajith for your constant love, care and guidance throughout my whole life. I thank my friends who have become my family in New Zealand, Nishadi, Vishan, Neranjala, Roshan, Pamiru, Mindi, Ravindu, Sampath, Chathurani, Nipuni, Waruna and Indika. I also thank Gayani and Ruwanthi for always putting a smile on my face. I thank my dearest friend Anurika, for our encouraging and joyful conversations and my sweetest nieces, nephews and my cousins who entertain me from 7000 miles away and finally my ever-loving boyfriend Kasun for your love and patience throughout this incredible journey. I thank all of you from the bottom of my heart, I am forever grateful! v Biology and behaviour of New Zealand ladybirds vi Biology and behaviour of New Zealand ladybirds Thesis abstract .......................................................................................................................................
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