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Uvic Thesis Template Influence of Seasonally Variable Hypoxia on Epibenthic Communities in a Coastal Ecosystem, British Columbia, Canada by Jackson Wing Four Chu B.Sc., Simon Fraser University, 2006 M.Sc., University of Alberta, 2010 A Dissertation Submitted in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY in the Department of Biology Jackson Wing Four Chu, 2016 University of Victoria All rights reserved. This dissertation may not be reproduced in whole or in part, by photocopy or other means, without the permission of the author. ii Supervisory Committee Influence of Seasonally Variable Hypoxia on Epibenthic Communities in a Coastal Ecosystem, British Columbia, Canada by Jackson Wing Four Chu B.Sc., Simon Fraser University, 2006 M.Sc., University of Alberta, 2010 Supervisory Committee Dr. Verena Tunnicliffe (Department of Biology, School of Earth and Ocean Sciences) Supervisor Dr. Francis Juanes (Department of Biology) Departmental Member Dr. Rana El-Sabaawi (Department of Biology) Departmental Member Dr. Roberta Hamme (School of Earth and Ocean Sciences) Outside Member iii Abstract Supervisory Committee Dr. Verena Tunnicliffe (Department of Biology, School of Earth and Ocean Sciences) Supervisor Dr. Francis Juanes (Department of Biology) Departmental Member Dr. Rana El-Sabaawi (Department of Biology) Departmental Member Dr. Roberta Hamme (School of Earth and Ocean Sciences) Outside Member Natural cycles of environmental variability and long-term deoxygenation in the ocean impose oxygen deficiency (hypoxia) on marine communities. My research exploits a naturally occurring hypoxia cycle in Saanich Inlet, British Columbia, Canada where I combined spatial surveys with remotely operated vehicles, ecological time-series from the subsea cabled observatory VENUS, and lab-based respirometry experiments to examine the influence of seasonally variable oxygen conditions on epibenthic communities. In situ oxygen thresholds established for dozens of fish and invertebrate species in this system show they naturally occur in lower oxygen levels than what general lethal and sublethal thresholds would predict. Expansion of hypoxic waters induced a loss of community structure which was previously characterized by disjunct distributions among species. Communities in variable hypoxia also have scale-dependent structure across a range of time scales but are primarily synchronized to a seasonal oscillation between two phases. Time-series revealed timing of diurnal movement in the slender sole Lyopsetta exilis and reproductive behavior of squat lobster Munida quadrispina in the hypoxia cycle. Hypoxia-induced mortality of sessile species slowed the rate of community recovery after deoxygenation. The 10-year oxygen time-series from VENUS, revealed a significant increase in the annual low-oxygen period in Saanich Inlet and that deoxygenation has occurred in this system since 2006. Differences in the critical oxygen crit thresholds (O2 ) and standard metabolic rates of key species (spot prawn Pandalus platyceros, slender sole, and squat lobster) determined the lowest in situ oxygen at which populations occurred and explained disproportionate shifts in distributions and iv crit community respiration. Finally, a meta-analysis on global O2 reported for crustaceans showed that hypoxia tolerance differs among major ocean basins. Long-term trends of deoxygenation suggest a future regime shift may occur when the duration at which a system remains below critical oxygen levels exceeds the time needed for communities to recover. Species-specific traits will determine the critical threshold and the nature of the community response in systems influenced by variable states of oxygen deficiency. However, oceanographic and evolutionary history provides context when determining the regional response of benthic communities influenced by rapidly changing environments. v Table of Contents Supervisory Committee ...................................................................................................... ii Abstract .............................................................................................................................. iii Table of Contents ................................................................................................................ v List of Tables ................................................................................................................... viii List of Figures .................................................................................................................... ix Acknowledgments............................................................................................................... x Chapter 1: General Introduction ......................................................................................... 1 Background ..................................................................................................................... 1 Oxygen loss in the ocean ............................................................................................ 1 Circulation of oxygen in the northern Pacific Ocean .................................................. 1 Multi-level response of metazoan life to low oxygen ................................................. 2 Seasonal hypoxia cycle in Saanich Inlet ..................................................................... 6 Research objectives ..................................................................................................... 8 Methodological approach.......................................................................................... 11 Literature cited .............................................................................................................. 12 Chapter 2: Oxygen limitations on marine animal distributions and the collapse of epibenthic community structure during shoaling hypoxia ................................................ 16 Preface........................................................................................................................... 16 Abstract ......................................................................................................................... 16 Introduction ................................................................................................................... 17 Materials and methods .................................................................................................. 21 Study site ................................................................................................................... 21 Benthic ROV transects .............................................................................................. 23 Video analysis and data management ....................................................................... 24 In situ oxygen limits for marine taxa ........................................................................ 25 Community-level organization by oxygen gradients ................................................ 27 Long-term oxygen profile from VENUS .................................................................. 28 Results ........................................................................................................................... 29 Seafloor oxygen profiles ........................................................................................... 31 The epibenthic animal community............................................................................ 32 Community-level reorganization during the 2013 hypoxia cycle ............................. 35 Long-term oxygen profile from VENUS .................................................................. 42 Discussion ..................................................................................................................... 43 Hypoxia thresholds in the northeast Pacific Ocean .................................................. 45 Hypoxia in the northeast Pacific Ocean .................................................................... 46 Community responses to hypoxia expansion ............................................................ 48 Acknowledgements ....................................................................................................... 50 Literature cited .............................................................................................................. 51 Chapter 3: Scale-dependent processes influence the temporal structure of epibenthic communities in a seasonally hypoxic system ................................................................... 60 Preface........................................................................................................................... 60 Introduction ................................................................................................................... 60 vi Materials and methods .................................................................................................. 63 Saanich Inlet.............................................................................................................. 63 Ocean Networks Canada camera systems ................................................................. 65 Imagery processing and data management ............................................................... 68 Statistical Analyses ................................................................................................... 69 Temporal changes in diversity .................................................................................
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