I HOW IS a FAMILY of SEDENTARY MARINE
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HOW IS A FAMILY OF SEDENTARY MARINE FISHES SHAPED BY ITS HABITATS, PREY, AND PREDATORS? by Clayton Garin Manning B.Sc. (First-Class Hons), University of Calgary, 2012 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE In THE FACULTY OF GRADUATE AND POSTDOCTORAL STUDIES (Zoology) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) August 2017 ©Clayton Garin Manning, 2017 i Abstract Overall, this thesis expands on our ecological understanding of a group of biologically diverse marine fishes by investigating how they are shaped by their habitats, prey, and predators. In my first data chapter, I used the seahorse Hippocampus whitei as a case study for investigating the ecological correlates of syngnathid abundance and distributions. Expanding on research that had looked at how either their habitats, prey, or predators affected their populations, I considered all three components in a single holistic approach. I investigated these correlations at two scales: among different seagrass beds (200-6000 m apart), and within a single seagrass bed (<100 m in size). I found that habitat, prey, and predator variables all correlated with seahorse density or height distributions to varying extents, depending on the scale of the study. Total predators was negatively associated with seahorse density across seagrass beds, the only ecological variable that was correlated with seahorses across beds. Within seagrass beds, seahorse locations correlated with greater depth, denser seagrass, more prey types, and fewer predators. In my second data chapter, I reviewed the diets and feeding behaviour of syngnathids, bringing together, summarizing, and providing new insights on a large amount of fragmented information on the topic. I answered three central questions. 1. How do syngnathids eat? 2. How does feeding and diet vary across a morphogically diverse family of fishes? 3. How does feeding and diet vary across a family of fishes that lives in a three- dimensional space? I answered question 1 by summarizing a number of different studies on the morphologies and kinematics of syngnathid feeding events. I answered questions 2 and 3 using a meta-analysis on syngnathid diets found the literature. Overall, I found there to be a large amount of variation in syngnathid diets that I hypothesize is caused by large ii differences in prey availability. Of the explained variation, I found their diets were most strongly correlated with their relative snout lengths and gape sizes. These feeding morphologies also had high phylogenetic signal, suggesting that dietary differences across genera were largely explained by how they differed with respect to these morphologies. iii Lay Summary Overall, this thesis investigates how syngnathids—a family of fish that includes seahorses and pipefishes—are affected by their habitats, prey, and predators. First, I looked at how seahorse abundance was affected by habitat, prey, and predators in eastern Australia. I found that each of these variables affected seahorse density or distributions to varying extents, depending on the scale of the study. Next, I reviewed syngnathid feeding, and provided a detailed summary on how syngnathids eat, and also analyzed what they eat. Overall, I found a lot of variation in what syngnathids eat, and suspect this is because there is a lot of variation in what is available to them in their environments. I also found their diets were best explained by the size and shapes of their snouts—highly advanced body parts that have evolved to help these fish feed on fast prey. iv Preface The research questions and methodological design of my thesis were developed in collaboration with my co-supervisors, Drs. Amanda Vincent and Sarah Foster. I collected all of the data used for Chapter 2 with the help of my host supervisor in Australia Dr. Dave Harasti, and my research assistants. I collected the data and information used in Chapter 3 from peer-reviewed and grey literature. I carried out all analyses, and prepared all manuscripts in this thesis, with substantial input from Amanda Vincent and Sarah Foster. A version of Chapter 2 is in the final stages of preparation for publication. As per our target journal’s requirements, the writing is in passive voice. I am the lead author, along with my co-supervisors Drs. Amanda Vincent and Sarah Foster, and my host-supervisor Dr. Dave Harasti, who provided guidance, logistical support, and assisted with fieldwork. I conducted all of the research for Chapter 2, and wrote the paper in collaboration with Drs. Sarah Foster and Amanda Vincent. Alistair Poore (University of New South Wales) helped train my assistants and me in crustacean identification, helped mold the study design, and provided statistical help. Meagan Abele and Natalie Scadden, my research assistants, were vital to the data collection and safety management of my project. A version of Chapter 3 is in the final stages of preparation for publication. I am the lead author, along with my co-supervisors Drs. Amanda Vincent and Sarah Foster. I conducted the analysis for Chapter 3 and wrote the paper with Drs. Sarah Foster and Amanda Vincent. I collected and tabulated the data used for Chapter 3 from primary and grey literature. v All field research undertaken in Chapter 2 was done in accordance with the University of British Columbia's Animal Care Committee permit A12-0288 and the NSW DPI Animal Care and Ethics Committee permit 15/01. vi Table of Contents Abstract .................................................................................................................................................... ii Lay Summary ......................................................................................................................................... iv Preface ...................................................................................................................................................... v Table of Contents................................................................................................................................vii List of Tables ......................................................................................................................................... xi List of Figures ...................................................................................................................................... xiv Acknowledgements ......................................................................................................................... xvii Chapter 1 Introduction ....................................................................................................................... 1 1.1 Rationale ................................................................................................................................ 1 1.2 Background ........................................................................................................................... 1 1.3 Research objectives and thesis outline ........................................................................ 6 Chapter 2 Ecological correlates of White's seahorse (Hippocampus whitei) abundance and size distributions at different spatial scales ....................................................................... 8 2.1 Introduction .......................................................................................................................... 8 2.2 Materials & Methods ....................................................................................................... 13 2.2.1 Study species ................................................................................................................................... 13 2.2.2 Study locations & design ............................................................................................................ 13 2.2.3 Seahorse surveys ........................................................................................................................... 17 2.2.4 Predator surveys ........................................................................................................................... 18 2.2.5 Seagrass surveys ............................................................................................................................ 19 vii 2.2.6 Prey surveys .................................................................................................................................... 19 2.2.7 Predicting covariates within Little Beach ......................................................................... 20 2.2.7.1 Seagrass and prey ................................................................................................................... 20 2.2.7.2 Predators .................................................................................................................................... 21 2.2.8 Statistical analyses ........................................................................................................................ 22 2.3 Results ................................................................................................................................ 25 2.3.1 Study scale: among all seagrass beds ................................................................................. 25 2.3.1.1 Seahorse survey summary statistics .............................................................................. 25 2.3.1.2 Correlates of seahorse density & seahorse height ...................................................