The Role of Boldness and Other Personality Traits in the Ecology of Juvenile Marine Fishes
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ResearchOnline@JCU This file is part of the following reference: White, James Ryan (2015) The role of boldness and other personality traits in the ecology of juvenile marine fishes. PhD thesis, James Cook University. Access to this file is available from: http://researchonline.jcu.edu.au/45988/ 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] and quote http://researchonline.jcu.edu.au/45988/ The role of boldness and other personality traits in the ecology of juvenile marine fishes Thesis submitted by James Ryan White BSc (University of California, Santa Cruz) MAppSci (James Cook University) For the degree of Doctor of Philosophy College of Marine and Environmental Sciences James Cook University Townsville, Queensland November 2015 i Statement of the Contribution of Others Financial Support James Cook University International Postgraduate Scholarship College of Marine and Environmental Sciences ARC Centre of Excellence for Coral Reef Studies AIMS@JCU Statistical and Analytical Support Prof. Mark McCormick Dr. Maud Ferrari Dr. Rabiul Beg Dr. Niels Dingemanse Dr. Ned Dochtermann Supervision Prof. Mark McCormick Dr. Mark Meekan Editorial Support Prof. Mark McCormick Dr. Mark Meekan ii Declaration on Ethics This research presented and reported in this thesis was conducted in compliance with the National Health and Medical Research Council (NHMRC) Australian Code of Practice for the Care and Use of Animals for Scientific Purposes, 7th Edition, 2004 and the Queensland Animal Care and Protection Act, 2001. The proposed research study received animal ethics approval from the JCU Animal Ethics Committee Approval Numbers # A1067 & A1720. iii Acknowledgements Numerous people have provided help and support throughout my thesis. First, I would like to thank my primary supervisor, Mark McCormick. Mark was always quick to return drafts of my papers, with useful comments and insights with larger ecological perspective. He always allowed me the independence I needed to work freely and supported my numerous trips to Lizard Island for my own work and for collaborations with his contacts. My secondary supervisor, Mark Meekan, provided very thorough edits on my manuscripts and constant direct support in the field. Mark was always available to provide life and career advice, as well as help me drink any beer that would otherwise have been left behind in the field. I would also like to thank Maud Ferrari and Shaun Killen for their guidance with statistical approaches and experimental design, as well as a lot of great laughs and memories at Lizard Island. I’ve had a lot of great volunteers in the field over the course of my thesis. Taylor Bodine, Caterina Schlott, Miwa Takahashi, Dylan Simonson, Marta Bertrand, Jon Smart and Kate Schoenrock all dedicated their time, effort and sanity working with me over long days at Lizard. Thanks guys for putting up with my sense of humor and tests of your patience! I would not have gotten through the trials and tribulations of my thesis without the emotional support and laughter provided by my friends over the years. Rohan Brooker, Bridie Allan, Cecilia Villacorta Rath, Matt Mitchell, Ian McLeod, Jackie Davies, Jennifer Atherton, Jon Smart, Luke Pedini, Matt Jankowski, Melanie Trapon, Rahel Zemoi, Richard Duffy, Samantha Munroe (my zombie-apocalypse partner), Tom Roberts, Kate Schoenrock, Daniel Cooper, Stephen Staples, Jennifer Hodge, Scott Harte, Crystal Neligh, Daniel Dixson, Donald Warren, Eva McClure, Terry Efird, Moorea Efird, Chris Reeves and Tory Chase all helped make my time here wonderful and memorable and supported me from near and afar. iv Ceci and Matt were especially supportive and valuable during my early years. They will be lifelong friends. Bridie single-handedly altered my perspective on the capabilities of human beings. I watched her grow from a relatively self-conscious, doubtful and inexperienced field scientist to a BAMF who kicks ass every day. Somehow she manages to be a full-time mom of two, wife, cook, PhD student, part-time research assistant and super-hero all at the same time. Knowing her makes me want to be a better person and shows me what we are all truly capable of with hard work and dedication. Bridie, your friendship made my time here worthwhile and is something I will always cherish. My lab mates have always been a source of amusement and support. Thanks to Katy Korkill, Lucy Holmes, Ryan Ramasamy, Govinda Liénart, Justin Rizzari, Davina Poulos, Maria Palacios and Lauren Nadler. Also, a special thanks to the staff at Lizard Island Research Station; Anne Hogget, Lyle Vail, Maryanne Pearce and Lance Pearce. They made Lizard Island such an easy, well- designed, and wonderful place to work. I would like to acknowledge the financial support from James Cook University, AIMS@JCU, the ARC CoE, and the College of Marine and Environmental Science. Without their support, my work wouldn’t have been possible. Finally, I would like to thank my family. My beautiful mother Susie, thank you for believing me and supporting my dreams in whatever way you could. Tim, you taught me what it means to be a strong man who supports his family. To Jeff, you are more than my twin brother, you are a part of me and I am a part of you. To Tyler, Troy, Beth and Savannah, when you guys showed up it changed my life dramatically, suddenly, and for the better. Watching you all grow up has taught me how to love unconditionally. And last but definitely not least, I would like to thank my Aunt Gail. She taught me to love the ocean, travelling, good food, and life in general. She has always been a personal v cheerleader, guidance counselor, financial supporter and advocate in my continued education and life goals. I would not be the man I am today without your love and encouragement. vi Abstract The theory of animal personality focuses on quantifying variation in behavior within and among individual organisms and attempts to account for the maintenance of differences in behavior that occur in a consistent manner among individuals. Personality has potentially important ecological consequences (e.g. behavioral tradeoffs) and can be shaped by population dynamics through selective mortality. Flexibility in behavior is advantageous for organisms that transition between stages of a complex life history. However, various constraints can set limits on plasticity, giving rise to the existence of personalities that have associated costs and benefits. One particularly important behavioral trait, boldness, is defined as the propensity of an animal to engage in risky behavior. Many variations of novel-object or novel-environment tests have been used to quantify the boldness of animals, although the relationship between test outcomes has rarely been investigated. Furthermore, the relationship of boldness measures to any ecological aspect of fitness is generally assumed, rather than measured directly. Understanding the costs and benefits of different behavioral phenotypes requires a greater understanding of structure and temporal consistency of intra-individual behaviors. More research is necessary for identifying the traits with potential fitness costs or showing how any trade-offs are manifested. This study therefore investigated the situational and temporal consistency of behavior, appropriateness of various boldness measures, and the relationships between different behavioral traits in order to better understand how coral reef fishes balance trade-offs related to risk. To understand the stability of fish behavior across various field and laboratory settings, there is a need to understand the behavioral structure throughout different situations. Chapter 2 tested for any evidence in consistency of behavior across situations in juveniles of a common damselfish, Pomacentrus amboinensis (Pomacentridae) at the transition between larval habitats in the plankton and juvenile habitats on the reef. vii Naïve fish leaving the pelagic phase to settle on reefs were caught by light traps and their behaviors observed using similar methods across three different situations (small aquaria, large aquaria, field setting); all of which represent low risk and well-sheltered environments. Seven behavioral traits were compared within and among individuals across situations to determine if consistent behavioral syndromes existed. No consistency was found in any single or combination of behavioral traits for individuals across all situations. We suggest that high behavioral flexibility is likely beneficial for newly-settled fishes at this ontogenetic transition and it is possible that consistent behavioral syndromes are unlikely to emerge in juveniles until environmental experience is gained or certain combinations of behaviors are favored by selective mortality. Despite the lack of evidence for behavioral syndromes, individual juvenile coral reef fish are likely to show behavioral repeatability within a single situation, over time (i.e. personality). Chapter 3 documented a field and laboratory experiment that examined the consistency of measures of boldness, activity, and aggressive behavior in young P. amboinensis immediately following their transition between pelagic larval and benthic juvenile habitats. Newly-settled fish were observed in aquaria and in the field on replicated patches of natural habitat cleared of resident fishes.