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Investigations Into the Skeletal Mineralogy of Temperate and Polar Bryozoans

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Investigations Into the Skeletal Mineralogy of Temperate and Polar Bryozoans INVESTIGATIONS INTO THE SKELETAL MINERALOGY OF TEMPERATE AND POLAR BRYOZOANS JENNIFER L LOXTON, BSC MSC SUBMITTED FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN MARINE BIOLOGY TO HERIOT WATT UNIVERSITY, SCHOOL OF LIFE SCIENCES MARCH, 2014 The copyright in this thesis is owned by the author. Any quotation from the thesis or use of any of the information contained in it must acknowledge this thesis as the source of the quotation or information. Abstract This study aims to explore the skeletal mineralogy of temperate and Polar bryozoans, investigating variability within and between species in relation to methodological, biological and environmental factors. Oceans are becoming warmer and more acidic and it is becoming increasingly important to increase our knowledge about the responses of marine calcifiers to environmental conditions. Bryozoans are important components of the benthic community globally, however prior to this study relatively little was known about their skeleton composition. This study has contributed over 1700 new mineralogical analyses to the field and provided new skeletal profiles for 115 bryozoan species. The study represents by far the most comprehensive regional profile of bryozoan mineralogy to date with 79% of Scottish species analysed. Targeted experiments have resulted in the documentation of impacts of curational procedures on skeletal mineralogy, resulting in recommendations which are pertinent across taxa in the wider fields of both curation and palaeoclimatography. Evidence is presented of mineral localization in specific skeletal features in bryozoan skeletons which adds to the growing body of data showing MgCO3 localization for mechanical and ecological advantage in marine invertebrates. Prior to this study it was proposed that both “active” biological and “passive” environmental controls influence bryozoan skeletal mineralogy. Through the examination of both Polar and Temperate species, this study has provided evidence of biological control of bryozoan mineralogy, while finding no evidence of passive environmental control. This finding precludes the use of bryozoan mineralogy for palaeoclimatic interpretation, for the species included in this study, and it is recommended that future species are chosen carefully and thoroughly calibrated prior to their use as palaeothermometers. Further investigation into the effects of ecological specification on the temperature/mineralogy response may, however, prove an area for fruitful research, enabling prediction of climate change effects on the bryozoan skeletons of ecologically specialised species and providing insights into future changes in community composition. Acknowledgements The author would like to acknowledge her supervisors: Associate Professor Joanne S Porter; Associate Professor Piotr Kuklinski; Mary Spencer Jones; Doctor Philip Cowie and Professor James M Mair. Without the valuable guidance, support and intellectual contributions of this supervisory group, the study would not have been possible. The author would also like to thank the Marine Alliance for Science and Technology in Scotland (MASTS) and the Marine Environmental Research Group (MERG) at Heriot-Watt University for supporting and funding this Ph.D. Additional thanks are extended to the NERC Collaborative Gearing Scheme, Heriot-Watt Alumni Fund, and Catlin Group Limited for financing or sponsoring this project. Special thanks are extended to the Heriot- Watt Scientific Dive Team and the staff at the British Antarctic Survey and Rothera Base for invaluable field support. Table of Contents Chapter 1. Introduction ...................................................................................................... 1 1.1 Calcium carbonate in the oceans ............................................................................. 1 1.1.1 Marine calcifiers............................................................................................... 1 1.1.2 The oceans carbonate system ........................................................................... 1 1.2 Ocean chemistry and climate change ...................................................................... 3 1.2.1 Ocean acidification........................................................................................... 3 1.2.2 Ocean warming ................................................................................................ 4 1.2.3 Additional pressures ......................................................................................... 5 1.3 Calcium carbonate variation .................................................................................... 5 1.3.1 Mechanisms of calcium carbonate variation .................................................... 5 1.3.2 Understanding the past, establishing a baseline for the present and predicting the future ........................................................................................................................ 7 1.4 The Bryozoa ............................................................................................................ 8 1.4.1 Introducing the Bryozoa ................................................................................... 8 1.4.2 History of mineralogical analysis and techniques applied to bryozoans ....... 11 1.4.3 Mineralogical variation in Bryozoa ............................................................... 13 1.4.4 Known knowledge gaps and areas for advancement ..................................... 15 1.5 Aims and objectives .............................................................................................. 16 Chapter 2. The forgotten variables: The impact of common cleaning and preservation techniques on skeletal chemistry. ......................................................................................... 18 2.1 Abstract …………………………………………………………………………..18 2.2 General Introduction .............................................................................................. 19 2.2.1 Factors known to affect the mineralogy of calcium carbonate ...................... 19 2.2.2 Preservation of calcified organisms ............................................................... 20 2.2.3 Cleaning and preparation ............................................................................... 24 ii 2.3 Aim and Objectives ............................................................................................... 27 2.3.1 Background to purpose of study .................................................................... 27 2.3.2 Objectives of study ......................................................................................... 27 2.4 Materials and Methods .......................................................................................... 28 2.4.1 Experiment to investigate the effects of cleaning methods on Mg-calcite .... 28 2.4.2 Experiment to investigate the effects of preservation on Mg-calcite………..30 2.4.3 XRD analysis ................................................................................................. 31 2.4.4 CHN analysis ................................................................................................. 34 2.4.5 Inductively Coupled Plasma Atomic Emission Spectroscopy (ICP-AES) .... 35 2.4.6 Review of past cleaning and preservation methods ....................................... 35 2.4.7 Data analysis .................................................................................................. 35 2.5 Results ................................................................................................................... 37 2.5.1 Cleaning Methods ............................................................................................... 37 2.5.2 Preservation .................................................................................................... 47 2.6 Discussion ............................................................................................................. 52 2.6.1 The effect of cleaning on bryozoan mass and skeletal Mg-calcite ................ 52 2.6.2 The effect of preservation on bryozoan skeletal Mg-calcite .......................... 57 2.7 Summary and Conclusions .................................................................................... 65 Chapter 3. Mineralogy of Scottish Bryozoans ................................................................. 66 3.1 Abstract ................................................................................................................. 66 3.2 Introduction ........................................................................................................... 67 3.2.1 Bryozoan studies in Scotland ......................................................................... 67 3.2.2 Regional studies of mineralogy ...................................................................... 68 3.2.3 Why a regional study in Scotland?................................................................. 68 3.2.4 Previous mineralogical analyses of Scottish species ..................................... 69 3.3 Aims and Objectives ............................................................................................. 69 iii 3.3.1 Summary of purpose of study ........................................................................ 69 3.3.2 Objectives of study ......................................................................................... 69 3.4 Materials and Methods .......................................................................................... 70 3.4.1 Sample
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