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Diversity and Distribution of Heterotrophic Flagellates in the Arctic Ocean

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Diversity and Distribution of Heterotrophic Flagellates in the Arctic Ocean Diversity and distribution of heterotrophic flagellates in the Arctic Ocean Thèse Mary Thaler Doctorat en Océanographie Philosophiae Doctor (PhD.) Québec, Canada © Mary Thaler, 2014 2 Résumé Dans les environnements marins arctiques, les protistes unicellulaires constituent les premiers maillons du réseau trophique. Les flagellés hétérotrophes (HF) jouent un rôle clé au sein de ce réseau trophique comme brouteurs de bactéries et de phytoplanctons, étant broutés à leur tour par les microzooplanctons comme les dinoflagellés ou les ciliés. Les scientifiques prévoient que les changements environnementaux extrêmes qui ont présentement lieu dans l‟Océan Arctique transformeront ces communautés de protistes. Le sujet de cette thèse porte sur la composition taxonomique des communautés marines HF dans l‟Océan Arctique, et leur réponse aux facteurs environnementaux. L‟approche a été d‟utiliser le comptage sur microscope à l‟aide de l‟hybridation fluorescente in situ pour évaluer l‟abondance de deux taxons HF importants, le genre Cryothecomonas et le clade MAST-1 parmi les straménopiles marins. Une approche complémentaire a été de décrire la répartition de tous les taxons HF, y compris l‟ordre Cryomonadida, les straménopiles marins, Picozoa, Telonemia, et les choanoflagellés, par l‟utilisation du séquençage à haut débit. Les résultats des deux approches nous ont permis de capturer les tendances environnementales sur une large échelle géographique dans l‟Arctique. Il a été mis en évidence une composition taxonomique structurée, principalement dû à l‟influence de la glace de mer et d‟autres facteurs environnementaux. Les cellules de Cryothecomonas semblaient provenir de la glace de mer, et au sein de la colonne d‟eau ils se trouvaient les plus nombreuses près du bord de glace. Par contre, les trois sous-clades de MAST-1 étaient retrouvés principalement dans la colonne d‟eau, mais répartis différemment par rapport au couvert de glace et les zones marginales de glace. La composition de la totalité de la communauté HF variait aussi par région, avec une plus grande importance de Telonemia et des choanoflagellés dans le Bassin du Canada. Pour plusieurs taxons, nous avons pu identifier un ou deux phylotypes dominants pour une région donnée. L‟importance relative de ces taxons devrait changer lors de la retraite de glace continue dans l‟Arctique, menant à des changements importants dans les réseaux trophiques et les cycles biogéochimiques iii Abstract In marine environments, single-celled protists form the initial links of the food web. Heterotrophic flagellates (HF) play a key role by grazing on bacteria and phytoplankton, being grazed upon in their turn by microzooplankton such as dinoflagellates and ciliates. The extreme environmental changes currently taking place in the Arctic Ocean are expected to transform protist communities. The subject of this thesis is the taxonomic composition of marine HF communities in the Arctic Ocean, and their response to environmental factors. The approach was to use fluorescent oligonucleotide probes to assess the abundance of two important HF taxa, the genus Cryothecomonas and the clade MAST-1 of the marine stramenopiles, via microscope counts. A complementary approach was to describe distribution of all HF taxa, including Cryomonadida, marine stramenopiles, Picozoa, Telonemia and choanoflagellates, by means of high-throughput sequencing. Results from these two approaches allowed us to capture broad environmental trends over a large geographic scale in the Arctic. A picture emerged of taxonomic composition largely structured by the influence of sea ice and other environmental factors. Cryothecomonas cells are inferred to live principally in the sea ice, and in the water column are more numerous close to the ice edge, whereas three sub-clades of MAST-1 are all found principally in the water column but are distributed differently relative to ice cover and marginal ice zones. The composition of the total HF community also varied by region, with a greater importance of Telonemia and choanoflagellates in the Canada Basin. For several taxa it was possible to identify one or two dominant phylotypes in a given region. The relative importance of these taxa is expected to change as sea ice retreat continues in the High Arctic, leading to important changes in trophic webs and biogeochemical cycles. v Table de Matières RÉSUMÉ ........................................................................................................................................................ III ABSTRACT ...................................................................................................................................................... V TABLE DE MATIÈRES .............................................................................................................................. VII LISTE DES TABLEAUX .............................................................................................................................. XI LISTE DES FIGURES ............................................................................................................................... XIII LISTE DE TABLEAUX SUPPLÉMENTAIRES .................................................................................... XVII LISTE DES ABRÉVIATIONS ...................................................................................................................XIX REMERCIEMENTS ...................................................................................................................................XXI AVANT-PROPOS .................................................................................................................................... XXIII CHAPTER 1 : INTRODUCTION GÉNÉRALE ............................................................................................. 1 1.01. INTRODUCTION ................................................................................................................................... 1 1.02. LES FLAGELLÉS HÉTÉROTROPHES ET LES RÉSEAUX TROPHIQUES MARINS ........................................... 1 1.03. L‘IMPORTANCE ÉCOLOGIQUE DE LA TAILLE DES ORGANISMES............................................................ 2 1.04. L‘EFFET DES CHANGEMENTS CLIMATIQUES SUR LES ORGANISMES DE L‘ARCTIQUE ............................ 2 1.05. LES OBJECTIFS DE CETTE THÈSE .......................................................................................................... 3 1.06. LES TAXONS HF .................................................................................................................................. 4 1.06.1. Cryomonadida ............................................................................................................................ 4 1.06.2. Choanoflagellés .......................................................................................................................... 4 1.06.3. Straménopiles Marins (MAST) ................................................................................................... 5 1.06.4. Picozoa ....................................................................................................................................... 5 1.06.5. Telonemia ................................................................................................................................... 5 1.07. MÉTHODOLOGIE ................................................................................................................................. 6 1.07.1. Hybridation fluorescente in situ (FISH) ..................................................................................... 6 1.07.2. Séquençage à haut-débit ............................................................................................................. 7 1.08. ANALYSES STATISTIQUES ................................................................................................................... 8 1.08.1. Les différents types de variables ................................................................................................. 8 1.08.2. Analyses de correspondance et de chemin.................................................................................. 8 1.08.3. Analyse canonique de correspondance ....................................................................................... 9 1.09. RÉGION D‘ÉTUDE .............................................................................................................................. 10 1.09.1. Nord de la Mer de Baffin .......................................................................................................... 10 1.09.2. Archipel Arctique Canadien ..................................................................................................... 10 1.09.3. Mer de Beaufort ........................................................................................................................ 11 1.09.4. Bassin du Canada ..................................................................................................................... 11 1.09.5. Mer des Tchouktches ................................................................................................................ 11 1.10. PLAN DE LA THÈSE ............................................................................................................................ 11 CHAPTER 2 : DISTRIBUTION AND DIVERSITY OF A PROTIST PREDATOR CRYOTHECOMONAS (CERCOZOA) IN ARCTIC MARINE WATERS .................................................... 21 RÉSUMÉ .......................................................................................................................................................
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