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Winners and Losers in a Changing Ocean: Impact on the Physiology and Life History of Pteropods in the Scotia Sea; Southern Ocean

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Winners and Losers in a Changing Ocean: Impact on the Physiology and Life History of Pteropods in the Scotia Sea; Southern Ocean Winners and losers in a changing ocean: Impact on the physiology and life history of pteropods in the Scotia Sea; Southern Ocean A thesis submitted to the School of Environmental Sciences of the University of East Anglia in partial fulfilment of the requirements for the degree of Doctor of Philosophy By Jessie Gardner June 2019 © This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with the author and that use of any information derived therefrom must be in accordance with current UK Copyright Law. In addition, any quotation or extract must include full attribution. Winners and losers in a changing ocean: Impact on the physiology and life history of pteropods in the Scotia Sea; Southern Ocean. © Copyright 2019 Jessie Gardner 2 Winners and losers in a changing ocean: Impact on the physiology and life history of pteropods in the Scotia Sea; Southern Ocean. Abstract The Scotia Sea (Southern Ocean) is a hotspot of biodiversity, however, it is one of the fastest warming regions in the world alongside one of the first to experience ocean acidification (OA). Thecosome (shelled) pteropods are planktonic gastropods which can dominate the Scotia Sea zooplankton community, form a key component of the polar pelagic food web and are important contributors to carbon and carbonate fluxes. Pteropods have been identified as sentinel species for OA, since their aragonitic shells are vulnerable to dissolution in waters undersaturated with respect to aragonite. In this thesis I investigate the impact of a changing ocean on the physiology and life history of pteropods in the Scotia Sea. Firstly, I culture early stage pteropods within OA and warming conditions predicted to occur in 2100 (Chapter 2). I demonstrate that larval shell morphology and survival rates are detrimentally affected in these conditions. Secondly, I constrain the life cycle and population dynamics of pteropods collected over a year from a sediment trap deployed on a moored platform (Chapter 3). I show that Limacina helicina and Limacina retroversa both have distinct life history strategies, although, spawning of both species corresponds to phytoplankton blooms. Thirdly, I establish a baseline vertical and biogeographical distribution of pteropods using historical samples (Chapter 4). I elucidate the geographical range edges of L. retroversa and L. helicina, as well as vertical migration patterns in relation to predation threat. Finally, I examine in-situ pteropod shell condition in relation to carbonate chemistry using net and oceanographic samples collected during two recent cruises (Chapter 5). I demonstrate that larval shells are susceptible to dissolution on exposure to aragonite undersaturation, however, later life stages display some resilience, since shell dissolution is confined to breaches in the periostracum. Overall, I recommend that continued monitoring, combined with bioassays and mesocosm work, will be essential in identifying the continued threat to pteropods from rapid environmental changes. 3 Winners and losers in a changing ocean: Impact on the physiology and life history of pteropods in the Scotia Sea; Southern Ocean. Contents Abstract ..................................................................................................................... 3 Contents .................................................................................................................... 4 List of Figures ............................................................................................................. 8 List of Tables ............................................................................................................ 18 Acknowledgements .................................................................................................. 20 1 Introduction ..................................................................................................... 21 1.1 Pteropod biology and role in the marine ecosystem ........................................ 22 1.2 Pteropod life history and physiology ................................................................ 26 1.3 Pteropods in a changing world: an overview of current experimental research 28 1.3.1 Ocean acidification .................................................................................... 28 1.3.2 Oceanic warming ...................................................................................... 37 1.3.3 Multiple drivers ......................................................................................... 38 1.4 An overview of in-situ pteropod conditions in a changing world ..................... 40 1.5 Study region ...................................................................................................... 40 1.6 Thesis aims and approach ................................................................................. 43 2 Southern Ocean pteropods at risk from ocean acidification and warming ........... 45 2.1 Introduction ...................................................................................................... 46 2.1.1 A changing world....................................................................................... 46 2.1.2 Early life history stages and oceanic change ............................................. 46 2.1.3 Southern Ocean pteropods ....................................................................... 47 2.1.4 Larval pteropods in a changing world ....................................................... 48 2.1.5 Chapter aims ............................................................................................. 48 2.2 Methods ............................................................................................................ 50 2.2.1 Pteropod collection and culturing ............................................................ 50 2.2.2 Experimental design .................................................................................. 52 2.2.3 Seawater manipulation ............................................................................. 55 2.2.4 Statistical analysis ..................................................................................... 57 2.3 Results ............................................................................................................... 58 2.3.1 Change in larval pteropod mortality ......................................................... 58 2.3.2 Change in larval pteropod shell size ......................................................... 59 2.3.3 Change in larval pteropod shell morphology ............................................ 60 2.3.4 Overall shell morphology .......................................................................... 63 2.4 Discussion .......................................................................................................... 64 2.4.1 Ocean acidification increases larval mortality .......................................... 64 4 Winners and losers in a changing ocean: Impact on the physiology and life history of pteropods in the Scotia Sea; Southern Ocean. 2.4.2 Ocean acidification and warming decreased shell size ............................. 65 2.4.3 Shell morphology is altered in a high CO2 world ....................................... 66 2.4.4 Concluding remarks and summary ........................................................... 68 3 Southern Ocean pteropod life history and population dynamics ........................ 70 3.1 Introduction ...................................................................................................... 71 3.1.1 Southern Ocean pteropod life cycle and population dynamics ................ 71 3.1.2 Life stage identification ............................................................................. 74 3.1.3 Sediment traps as tools for pteropod population studies ........................ 75 3.1.4 Sampling area ............................................................................................ 76 3.1.5 Chapter Aims ............................................................................................. 78 3.2 Materials and Methods ..................................................................................... 79 3.2.1 Sample collection ...................................................................................... 79 3.2.2 Trapping efficiency .................................................................................... 81 3.2.3 Sample processing .................................................................................... 83 3.2.4 Species identification and shell morphometrics ....................................... 84 3.2.5 Classification of shell condition ................................................................. 86 3.2.6 Cohort and group identification................................................................ 87 3.2.7 Shell growth, spawning behaviour and longevity ..................................... 88 3.2.8 Environmental parameters ....................................................................... 89 3.3 Results ............................................................................................................... 92 3.3.1 Species presence ....................................................................................... 92 3.3.2 Species and life stage abundance ............................................................. 93 3.3.3 Shell morphometrics ................................................................................. 96 3.3.4
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