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University of California, San Diego UNIVERSITY OF CALIFORNIA, SAN DIEGO Mesozooplankton responses to climate variability and contributions to microbial export A dissertation submitted in partial satisfaction of the requirements for the Degree Doctor of Philosophy in Oceanography by Bellineth Valencia Ramírez Committee in charge: Professor Michael R. Landry, Chair Professor Andrew E. Allen Professor Mark D. Ohman Professor Brian Palenik Professor Jonathan B. Shurin 2017 Copyright Bellineth Valencia Ramírez, 2017 All Rights Reserved The dissertation of Bellineth Valencia Ramírez is approved, and it is acceptable in quality and form for publication on microfilm and electronically: Chair University of California, San Diego 2017 iii TABLE OF CONTENTS SIGNATURE PAGE .................................................................................................................... iii TABLE OF CONTENTS .............................................................................................................. iv LIST OF FIGURES ..................................................................................................................... viii LIST OF TABLES ....................................................................................................................... xiii ACKNOWLEDGEMENTS ......................................................................................................... xvi VITA ............................................................................................................................................ xxi ABSTRACT OF THE DISSERTATION .................................................................................. xxiii INTRODUCTION .......................................................................................................................... 1 The biological pump and the role of mesozooplankton .......................................................... 1 Mesozooplankton in the oligotrophic North Pacific ............................................................... 3 Mesozooplankton in the California Current System ............................................................... 5 Dissertation outline ................................................................................................................. 9 References ............................................................................................................................. 12 CHAPTER 1 Environmental drivers of mesozooplankton biomass variability in the North Pacific Subtropical Gyre ........................................................................................................................... 22 Abstract ……………………………………………………………………………………. 23 Introduction …………………………..……………………………………………………. 23 Materials and methods ………………………….…………………………………………. 24 Sampling and Laboratory Analysis ……………………………………………………. 24 Long-Term Trend Analysis …………………………..………………………………... 24 Environmental Data …………………………..………………………………..………. 25 Mesozooplankton Biomass and Environmental Factors …………………………….… 25 Results …………………………..………………………………...………………………. 27 Temporal Variability in Mesozooplankton Biomass …………………...……………… 27 Monthly Mean Model …………………………..…………………………………...…. 27 Annual Mean Model …………………………..…………………………………….…. 28 Discussion …………………………..…………………………………………………..…. 31 Plankton Variability in the North Pacific Subtropical Gyre (NPSG) ……………….…. 31 Links to Climate Variability …………………………..…………………………….…. 32 Conclusions …………………………..…………………………………………………… 33 iv Acknowledgements ………………………….……………………………………………. 34 References …………………………..………………………………………………….…. 34 CHAPTER 2 Environmental effects on mesozooplankton size structure and export flux at station ALOHA, North Pacific Subtropical Gyre ..................................................................................... 37 Abstract ................................................................................................................................. 37 Introduction ........................................................................................................................... 38 Methods................................................................................................................................. 40 Sampling and Laboratory Analyses .................................................................................. 40 Biomass Size Structure ..................................................................................................... 41 Temporal Trends of Biomass ............................................................................................ 42 Carbon Cycling in the Euphotic Zone .............................................................................. 43 Active and Passive Export Fluxes..................................................................................... 44 Environmental Effects on Size Structure and Migrant Biomass ...................................... 45 Results ................................................................................................................................... 46 Biomass Size Structure ..................................................................................................... 46 Carbon Cycling in the Euphotic Zone .............................................................................. 47 Migrant Biomass and Active Export Flux ........................................................................ 48 Environmental Relationships ............................................................................................ 50 Discussion ............................................................................................................................. 51 Mesozooplankton Biomass Size Structure........................................................................ 51 Carbon Cycling in the Euphotic Zone .............................................................................. 54 Mesozooplankton and Export Flux ................................................................................... 56 Conclusions ........................................................................................................................... 60 Acknowledgments................................................................................................................. 61 References ............................................................................................................................. 70 CHAPTER 3 The role of salps and doliolids in feeding and export of protists during two anomalously warm years in the California Current System ......................................................... 80 Abstract ................................................................................................................................. 80 Introduction ........................................................................................................................... 81 Materials and methods .......................................................................................................... 83 Sample collection for metabarcoding ............................................................................... 83 Library construction and sequencing ................................................................................ 85 Bioinformatic analyses...................................................................................................... 86 v Synechococcus sequence analysis and classification ....................................................... 87 Data analyses .................................................................................................................... 87 Results ................................................................................................................................... 88 Sequence analyses ............................................................................................................. 88 General patterns of protists detected in four species of pelagic tunicates ........................ 89 Protists detected in guts and fecal pellets of pelagic tunicates ......................................... 90 Salpa aspera and Cyclosalpa affinis .................................................................................. 90 Doliolum denticulatum and Pegea socia ........................................................................... 92 Salp fecal pellets analyzed from sediment traps ............................................................... 93 Synechococcus strains ...................................................................................................... 93 Discussion ............................................................................................................................. 93 Major groups of protistan parasites................................................................................... 94 General patterns within and among tunicate species ........................................................ 95 Potential implications for protistan export ........................................................................ 98 Synechococcus strains .................................................................................................... 101 Conclusions ......................................................................................................................... 102 Acknowledgements ............................................................................................................. 103 References ..........................................................................................................................
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