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The Effects of Dissolved Organic Carbon on Pathways of Energy Flow, Resource Availability, and Consumer Biomass in Nutrient-Poor Boreal Lakes

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The Effects of Dissolved Organic Carbon on Pathways of Energy Flow, Resource Availability, and Consumer Biomass in Nutrient-Poor Boreal Lakes The effects of dissolved organic carbon on pathways of energy flow, resource availability, and consumer biomass in nutrient-poor boreal lakes By Joseph Tonin A thesis submitted to the Faculty of Graduate Studies of The University of Manitoba In partial fulfillment of the requirement of the degree of Master of Science Department of Entomology University of Manitoba Winnipeg, Manitoba Copyright © 2019 by Joseph Tonin i Abstract Over the past few decades, terrestrially derived dissolved organic carbon (DOC) has been recognized as a fundamental driver of food web productivity in nutrient poor lakes. The mechanisms that underlie these effects remain poorly understood, particularly for higher trophic levels including zooplankton, benthic invertebrates, and fish. In a survey of eight lakes in northwestern Ontario, I determined consumer biomass and used stable isotopes of carbon, nitrogen, and hydrogen to investigate relationships between DOC and pathways of energy flow, resource and habitat availability, and consumer biomass. Using Bayesian stable isotope mixing models, I found that hypolimnetic phytoplankton were an important resource for zooplankton in low-DOC lakes. With increased DOC concentrations, light attenuation increased and chlorophyll a concentrations below the thermocline were reduced relative to epilimnetic concentrations. At higher DOC concentrations, zooplankton acquired proportionately more energy from low quality terrestrial sources. Zooplankton biomass also declined with increasing utilization of terrestrial sources (allochthony), suggesting that terrestrial organic matter suppresses zooplankton productivity through simultaneous limitations on habitat and resource availability and quality. Based on biomass, the dominant fish species across my study lakes was White Sucker (Catostomus commersonnii). Bayesian mixing models indicated that allochthony by White Suckers increased with DOC and that greater allochthony was related to lower White Sucker biomass measured as catch-per-unit-effort (bCPUE). Both White Sucker bCPUE and chironomid biomass were positively related to mean light irradiance, with the highest biomasses of fish and chironomids occurring in lakes with a higher proportion of their volume in the photic zone. White Sucker bCPUE was strongly and positively correlated with chironomid biomass, ii suggesting that DOC-mediated resource limitation may influence fish productivity via reduced prey availability. iii Acknowledgments First, to Mike Paterson. I cannot thank you enough for providing me with this opportunity. I am grateful for everything you have done for me – from sending me to conferences and workshops to providing constructive comments and criticism. You always allowed me freedom in my work, but were always there to steer me in the right direction, and for that, I am thankful. Your passion for science rubs off on everyone you meet, and your patience and encouragement throughout this process has made me a better scientist. It has been a pleasure working with you, and I look forward to continuing our work together. Thank you to my committee members, Dr. Mike Rennie, and Dr. Neil Holliday for keeping this project on track. This thesis work is part of a large collaborative project Photons to Fish: Ecosystem Indicators of fish productivity (PHISH) – special thank you to all collaborators that have made this project such a success and a joy to be a part of. Specifically, Mike Paterson, Mike Rennie, and Scott Higgins, who were instrumental in getting this project off the ground. To the University of Waterloo, Rachel Henderson in particular, who supported everything isotope related. To Bryanna Sherbo, who was always there to provide field support and discuss the future of the project. Lastly, to Justin Budyk, thank you for your unrelenting positivity and hard-work in the field. Most importantly, thank you for being a great friend. This thesis is largely of my own work, but does rely on contributions from others, which are summarized in Table at the end of the acknowledgements. This project was supported by Manitoba Hydro, the IISD Experimental Lakes Area (IISD-ELA) graduate fellowship fund, Mitacs, and Fish Futures Inc. iv I was fortunate to be able to conduct my field-work at the world renowned IISD-ELA research facility. Not only did IISD-ELA provide a platform for me to conduct research, it provided me with an opportunity to explore and appreciate nature. I encourage anyone and everyone to consider donating to IISD-ELA to ensure it is well funded in perpetuity. The science conducted at IISD-ELA has a global impact, and in the face of large scale ecosystem change, there is a need for long-term monitoring programs and whole-ecosystem science to ensure the ongoing conservation and preservation of fresh water resources. Of course, part of what makes IISD-ELA such a fantastic place to do freshwater science is the people. Without the help of IISD-ELA staff, this project would not have been possible. Thank you to Lee Hrenchuk, Chandra Rodgers, Lauren Hayhurst, and Mike Rennie for always being there for anything and everything fish related. To Sonya Havens and Kelli-Nicole Croucher, thank you for always being accommodating to my sampling schedule and for generating high quality data. To Ken Sandilands and Paul Fafard, you have always been generous to me, whether lending me equipment or sending me data, and for that I thank you. Thank you to Mike Paterson, Scott Higgins, Vince Palace, and Craig Emmerton for always being there to talk about science – the conversations we had over a coffee break or a game of hoops were the source of inspiration for many of the ideas presented in this thesis or ideas I hope to explore in the future. Special thanks to operations staff, Roger Mollot and John Neal, for keeping camp running like a well-oiled machine, and to Jesse and Frank, who kept me well fed. Also, thank you to Stephen Paterson, who always made sure there was a bed available for me, the first aid room was well stocked, and for always saying yes to a canoe trip. In addition to staff, there are countless summer students to thank. IISD-ELA runs on the backs of students, and each and every one of you has contributed to making this thesis happen. v Whether it be collecting data for the long-term monitoring program, joining me on a canoe trip, or playing volleyball at the beach. My time in Winnipeg has been an enjoyable experience thanks in large part to the friendships I gained while working at IISD-ELA. Thank you all for making graduate school a little easier. To my family (Team Tonin) – Mom, Dad, John, Brett and Ben. You all have been so supportive throughout this process even though you all have no idea what it is I do. Lastly, I need to thank my partner, Ooma. This thesis is dedicated to you. Your love and encouragement has been a steadying presence in my life for the last six years, and without your support, I’m not sure I how I could have gotten through this. And for that, I thank you. vi Attribution of work done by J. Tonin and others in this thesis: Performed by J. Tonin Contributions from Others • Monthly collection of stable isotope samples • Analysis of stable isotope samples from all 8 study lakes in 2016 and 2017 (Environmental Isotope Laboratory, (water, seston, periphyton, zooplankton, University of Waterloo for carbon and benthic macroinvertebrates) nitrogen samples, and Colorado Plateau • Processing of stable isotope samples Stable Isotope Laboratory, Northern Arizona • Monthly collection of samples for University for hydrogen samples) zooplankton, eplimnetic chemistry, • Collection of zooplankton, chemistry, chlorophyll a, light, temperature and oxygen chlorophyll a, light, temperature and oxygen profiles from Lakes 164, 658 in 2016 and profiles from Lakes 223, 224, 239, 373, 442, 2017, as well as zooplankton in Lakes 442 626 (IISD-ELA Hydro-Lim field crew) and 373 in 2017 • Analysis of all water chemistry samples • Collection of benthic macroinvertebrate (IISD-ELA Chemistry Laboratory) samples from 5 lakes in 2016, and all lakes • Collection of fish from Lakes 223, 224, 239, in 2017 373 and 626 for biomass catch per unit effort • Processing, counting, and identification of all estimates (IISD-ELA Fish Crew); J. Tonin benthic macroinvertebrate samples assisted with collection in 2016, and for • Collection of fish from Lakes 164, 442, 658 L239 in 2017 for biomass catch per unit effort estimates • Counting and Identification of zooplankton • Collection of fish stable isotopes from 6 of 8 (Plankton R Us) lakes in 2017 • Chlorophyll a profiles (Bryanna Sherbo) • Analysis and interpretation of all data vii Table of Contents ABSTRACT ............................................................................................................................. ii ACKNOWLEDGMENTS ...................................................................................................... IV TABLE OF CONTENTS .................................................................................................... VIII LIST OF TABLES ................................................................................................................... X LIST OF FIGURES ............................................................................................................... XI CHAPTER 1: GENERAL INTRODUCTION AND LITERATURE REVIEW .................... 1 INTRODUCTION ...................................................................................................................................................1 BACKGROUND
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