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The Role of Sea Ice Algae-Produced Carbon in Arctic and Antarctic Food Webs Dependency of Polar Life on a Threatened Food Source

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The Role of Sea Ice Algae-Produced Carbon in Arctic and Antarctic Food Webs Dependency of Polar Life on a Threatened Food Source The role of sea ice algae-produced carbon in Arctic and Antarctic food webs Dependency of polar life on a threatened food source Dissertation with the aim of achieving a doctoral degree at the Faculty of Mathematics, Informatics and Natural Sciences Department of Biology of the University of Hamburg submitted by Doreen Kohlbach M. Sc. Biotechnology B. Eng. Biotechnology Hamburg 2017 The following evaluators recommend the admission of the dissertation: 1. Evaluator Prof. Angelika Brandt Universität Hamburg, Biozentrum Grindel (CeNak), Zoologisches Museum, Martin-Luther King-Platz 3, 20146 Hamburg 2. Evaluator Dr. Hauke Flores Alfred-Wegener-Institut Helmholtz-Zentrum für Polar-und Meeresforschung, Am Handelshafen 12, 27570 Bremerhaven i ii Table of contents Summary ............................................................................................................................................................. v Zusammenfassung ....................................................................................................................................... vii Abbreviations ................................................................................................................................................... ix 1. Introduction ................................................................................................................................................ 1 1.1 The Polar Oceans: Impact of environmental changes ........................................................... 1 1.2 Sea ice algae-produced carbon in polar ecosystems .............................................................. 6 1.3 Food webs in Polar Regions ............................................................................................................. 9 1.3.1 Arctic ...................................................................................................................................... 11 1.3.2 Antarctic ................................................................................................................................ 12 1.4 Biomarker concept ........................................................................................................................... 14 1.5 State-of-the-Art techniques in lipid and stable isotope analyses ................................... 15 1.6 Objectives of this thesis .................................................................................................................. 19 1.7 Publication outline ........................................................................................................................... 21 2. Thesis chapters …………………………………………………………………………………………….............. 2.1 Chapter I .................................................................................................................................... 25 The importance of ice algae-produced carbon in the central Arctic Ocean ecosystem: Food web relationships revealed by lipid and stable isotope analyses ................................... 2.2 Chapter II .................................................................................................................................. 57 Strong linkage of polar cod ( Boreogadus saida ) to sea ice algae-produced carbon: evidence from stomach content, fatty acid and stable isotope analyses ................................. 2.3 Chapter III .................................................................................................................................85 Ice algae-produced carbon ensures winter survival of young Antarctic krill Euphausia superba .............................................................................................................................................................. 2.4 Chapter IV .............................................................................................................................. 103 Spatio-temporal variability in the winter diet of larval and juvenile Antarctic krill (Euphausia superba ) in ice-covered waters ........................................................................................ 2.5 Chapter V ................................................................................................................................ 127 Overwintering of Weddell Sea under-ice community strongly linked to sea ice- associated food sources .............................................................................................................................. 2.6 Chapter VI ...........................................................................................................................................165 Euphausia superba , E. crystallorophias and Thysanoessa macrura in the Filchner Outflow System: Variability of carbon sources during summer assessed in a lipid and stable isotope study ..................................................................................................................................... 3. Discussion .................................................................................................................................. 193 3.1 Quantifying dietary contributions of ice algal produced carbon: Methodological aspects ........................................................................................................................................................193 3.2 Importance of ice algae-produced carbon to the Arctic food web ...............................197 3.2.1 Foundation of every food web study: The trophic baseline .................................198 iii 3.2.2 Consumption of ice algal carbon: Do season and region matter? ....................200 3.2.3 How big is the impact of the changing sea ice environment on top predators? .............................................................................................................................................205 3.3 Importance of ice algae-produced carbon to the Antarctic food web ........................210 3.3.1 Exceptional role in the Antarctic food web: Antarctic krill Euphausia superba 212 3.3.2 The unattended krill: Euphausia crystallorophias and Thysanoessa macrura ..213 3.4 Impact of ice algae as food source: Comparison between the hemispheres ...........215 4. Conclusion and Outlook ..................................................................................................................221 References .....................................................................................................................................................223 Acknowledgements ...................................................................................................................................259 Eidesstaatliche Erklärung ............................................................................................................ 261 iv Summary The underside of sea ice in Polar Regions represents a natural habitat for heterotrophic organisms, such as copepods and amphipods. This under-ice fauna plays a key role in transferring carbon synthesized by sea ice-associated (sympagic) microalgae into associated pelagic and benthic food webs of polar ecosystems. Animals at higher trophic levels are adapted to feed on the under-ice fauna as well as on pelagic zooplankton and nekton. Polar ecosystems thrive significantly on ice algae-produced carbon depending on different periods of the year. Thus, the under-ice fauna and the associated pelagic food web are largely affected by multi-scale climate changes accompanied by the reduction of sea ice coverage and an increasing duration of the melt season. Until now, however, the degree to which polar food webs depend on sea ice-derived carbon is unclear. The overall aim of this thesis is to quantify the transfer of ice algae-produced carbon from the sea ice into the under-ice community and from there into pelagic food webs in Arctic and Antarctic ecosystems, in order to improve our understanding of the potential ecological consequences of a changing sea ice environment for marine food web dynamics. Furthermore, spatial and seasonal differences in the utilization of ice algae- produced carbon within and between both hemispheres are investigated. The sample collection in the central Arctic Ocean was carried out during the RV ‘Polarstern ’ expedition ARK XXVII-3 (PS80, August-September 2012) within the Amundsen and Nansen Basins. In the Southern Ocean, samples were collected during the RV ‘ Polarstern ’ expeditions ANT XXIX-7 (PS81, August-October 2013) in the northern Weddell Sea and ANT XXIX-9 (PS82, December 2013-March 2014) offshore from the Filchner Ice Shelf. Trophic interactions of important representatives of Arctic and Antarctic food webs are studied using lipid fingerprinting, stable isotope analysis (SIA) of natural abundance bulk carbon and nitrogen (BSIA), and compound-specific SIA (CSIA) of fatty acids (FAs). From the distribution of algae-produced FAs in the consumers (= marker FAs), the origin of carbon produced by diatoms versus dinoflagellates in key Arctic species (Chapter I and II ) and key Antarctic species (Chapters III-VI ) is investigated. Stable isotope mixing models are used to quantify the relative contribution of bulk carbon and marker fatty acids derived from ice algae versus pelagic phytoplankton to the carbon budget of the organisms. Additionally, the stomach contents of polar cod Boreogadus saida (Chapter II ) and Antarctic krill Euphausia superba (Chapter IV ) are investigated to provide information on the most recent diet composition
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