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OFFSPRING CANNIBALISM and PARENTAL CARE in the PLAINFIN MIDSHIPMAN FISH Phd Thesis –Aneesh P

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OFFSPRING CANNIBALISM and PARENTAL CARE in the PLAINFIN MIDSHIPMAN FISH Phd Thesis –Aneesh P OFFSPRING CANNIBALISM AND PARENTAL CARE IN THE PLAINFIN MIDSHIPMAN FISH PhD Thesis –Aneesh P. H. Bose McMaster University – Psychology, Neuroscience and Behaviour OFFSPRING CANNIBALISM AND PARENTAL CARE IN THE PLAINFIN MIDSHIPMAN FISH BY ANEESH P. H. BOSE, B.Sc. A Thesis Submitted to the School of Graduate Studies in Partial Fulfilment of the Requirements for the Degree Doctor of Philosophy McMaster University © Aneesh Bose, July 2017 ii PhD Thesis –Aneesh P. H. Bose McMaster University – Psychology, Neuroscience and Behaviour DOCTOR OF PHILOSOPHY (2017) McMaster University (Psychology, Neuroscience & Behaviour) Hamilton, Ontario TITLE: Offspring cannibalism and parental care in the plainfin midshipman fish AUTHOR: Aneesh P. H. Bose, B.Sc. (University of Guelph) SUPERVISOR: Dr. Sigal Balshine, B.Sc. (University of Toronto), Ph.D. (Cambridge University) NUMBER OF PAGES: xiv, 270 iii PhD Thesis –Aneesh P. H. Bose McMaster University – Psychology, Neuroscience and Behaviour Thesis abstract Cannibalism of offspring is a common yet seemingly paradoxical phenomenon observed across a wide variety of taxa. Behavioural ecologists have been particularly interested in understanding this behaviour within the context of parental care. This is because, superficially, offspring cannibalism appears counterproductive to the goals of a caring parent, which are often achieved by nurturing and protecting their young. Despite the prevalence of offspring cannibalism in many taxa, we still have a poor evolutionary understanding of this intriguing behaviour. Many hypotheses have been proposed explaining why parents may terminate and consume offspring, however, empirical tests of these hypotheses frequently return mixed results, and few examples exist in which multiple hypotheses for cannibalism have been tested using the same model study system. Over the course of my PhD thesis, I tested different cannibalism hypotheses in a novel study organism, the plainfin midshipman fish (Porichthys notatus), a species of paternal care-giving toadfish that frequently engages in offspring cannibalism. I first characterized the factors that underlie male reproductive success in this species to provide a broader understanding of their reproductive ecology and a background for the cannibalism studies in my later chapters. I showed that both male size and nest size are important correlates of male reproductive success (e.g. mate attraction, egg acquisition, and rearing success) in this system, and that morphological, physiological, and ecological variables can also have an impact on reproductive success (Chapter 2). Next, I conducted a series of field and iv PhD Thesis –Aneesh P. H. Bose McMaster University – Psychology, Neuroscience and Behaviour laboratory studies to uncover the factors that select for offspring cannibalism in this species. In particular, I investigated whether offspring cannibalism serves to replenish dwindling energy reserves and/or occurs when paternity over a brood is likely to be low. I showed that although plainfin midshipman males endure a long and energetically taxing parental care period, they cannibalize offspring most frequently early in the breeding season, the time period when the males are in their best body condition but when male- male competition is also at its peak (Chapter 3). Using a direct comparison of males that had recently cannibalized offspring versus those that had not, I showed that the males with the lowest energy reserves were actually the least likely to have offspring in their digestive tracts, suggesting that offspring cannibalism is not driven by immediate energetic need (Chapter 4). I then tested whether males assess their paternity using direct or indirect offspring cues, and showed that males use the act of a nest take-over as reliable indirect cues of their paternity over a brood (Chapter 5). Altogether, my results further our understanding of the reproductive ecology of the plainfin midshipman fish, and provide a foundation upon which to assess and test between cannibalism hypotheses in this species and others. My thesis work demonstrates the advantage of studying multiple cannibalism hypotheses in a single study system, and through this approach we can gain a more accurate evolutionary understanding of how the phenomenon of offspring cannibalism is maintained in different species. v PhD Thesis –Aneesh P. H. Bose McMaster University – Psychology, Neuroscience and Behaviour Acknowledgements First and foremost, I would like to thank my supervisor, Dr. Sigal Balshine, for her mentorship during my PhD. I am very grateful to have had her as a supervisor, as her passion and enthusiasm for science has made me the scientist that I am today. I could not have picked a more encouraging and supportive lab for my graduate studies. I would also like to thank my supervisory committee, Dr. Jim Quinn and Dr. Reuven Dukas, for their advice and guidance. My work in the field would not have been possible without the help of a large group of fantastic collaborators on the west coast. Many thanks to Dr. Karen Cogliati for training me on “how to do field work”, and to Dr. Joe Sisneros for facilitating our work at my favourite field site, Dabob Bay in Washington, USA. A big thank you to Holly Howe, Henry Kou, Nick Luymes, Eric Balke, and Theresa Warriner for their hard labour collecting data with me in the intertidal zones. I am also extremely grateful to Pamela Walker and Ross Shepard, Captain Bill and Rubie Cogswell, Eileen Carr and family, Chuck and Sally Flader, and Michael and Dvori Balshine for their amazing hospitality while we conducted our field work. My lab work would also not have been possible without the help and generosity of Holly Hicklin, Dr. Henrik Kreiberg, Dr. Francis Juanes, Kieran Cox, Dr. Grant McClelland, Dr. Bryan Neff, and Malcolm Lau. A big thank you to the many past ABEL undergraduate students for their contributions to this body of work. And a huge thank you to Dr. Ben Bolker and Dr. Jonathan Dushoff for imparting a small portion of their statistical wizarding knowledge to me over the years. vi PhD Thesis –Aneesh P. H. Bose McMaster University – Psychology, Neuroscience and Behaviour Five years is a considerable duration and I’m grateful to have spent it with some great friends and lab mates, Susan Marsh-Rollo, Dr. Connie O’Connor, Dr. Cody Dey, Jess Miller, Dr. Laura Cirelli, Gates Imbeau, Dr. Brandon Paul, Dr. Fiona Manning, Tyler Pollock, Tammy Rosner, and Evan Borman. I would also like to thank my parents for their constant support and encouragement. Lastly, I am endlessly grateful to Dr. Erin McCallum for her love and support. I could not be more thankful to have her by my side while we face academia (and other paths in life) together. vii PhD Thesis –Aneesh P. H. Bose McMaster University – Psychology, Neuroscience and Behaviour Table of contents Thesis abstract ............................................................................................................................. iv Acknowledgements ..................................................................................................................... vi List of figures .............................................................................................................................. xi List of tables ................................................................................................................................ xi Declaration of academic achievement ........................................................................................ xii Chapter 1: General introduction ................................................................................................... 1 1.1 Theoretical background and motivation for thesis ................................................................. 1 1.2 Aims of the thesis .................................................................................................................. 6 1.3 Structure of the thesis .......................................................................................................... 14 1.4 References ........................................................................................................................... 20 Chapter 2: Male and resource quality as factors affecting male reproductive success in an intertidal fish ................................................................................................................................. 44 2.1 Abstract ............................................................................................................................... 44 2.2 Introduction ....................................................................................................................... 455 2.3 Methods ............................................................................................................................... 48 2.4 Results ................................................................................................................................. 57 2.5 Discussion ........................................................................................................................... 64 2.6 Acknowledgments ............................................................................................................... 71 2.7 References ........................................................................................................................... 72 2.8 Supplementary materials ....................................................................................................
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