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These De Doctorat De THESE DE DOCTORAT DE L'UNIVERSITE DE BRETAGNE OCCIDENTALE COMUE UNIVERSITE BRETAGNE LOIRE ECOLE DOCTORALE N° 598 Sciences de la Mer et du littoral Spécialité : Ecologie Marine Par Nicolas DJEGHRI Variability and Plasticity of the Nutrition of Zooxanthellate Jellyfishes Insights from experimental and field studies. Variabilité et Plasticité de la Nutrition des Méduses à Zooxanthelles Apports expérimentaux et de terrain. Thèse présentée et soutenue à Plouzané, le 2 décembre 2019 Unité de recherche : Lemar Rapporteurs avant soutenance : Composition du Jury : Cathy LUCAS Président du Jury Associate Professor, Frédéric JEAN National Oceanographic Centre, Royaume-Uni Professeur, Université de Bretagne Occidentale, France Fabrice NOT Directeur de Recherche CNRS, Cathy LUCAS Station Biologique de Roscoff, France Associate Professor, National Oceanographic Centre, Royaume-Uni Fabrice NOT Directeur de recherche CNRS, Station Biologique de Roscoff, France Gauthier SCHAAL Maitre de Conférences, Université de Bretagne Occidentale, France Pascal CLAQUIN Professeur, Université de Caen Normandie, France Delphine THIBAULT Maitre de Conférences, Institut Méditerranéen d’Océanologie, France Directeur de thèse Philippe PONDAVEN Maitre de Conférences, Université de Bretagne Occidentale, France Co-directeur de thèse Herwig STIBOR Professor, Ludwig-Maximilians-Universität München, Allemagne II Remerciements Je eu tout d’abord remercier mon directeur de thèse, Philippe Pondaven, et mon co- direteur Herig “tior. E preier lieu de ’aoir doé l’opportuité d’attauer ce sujet de thèse pour le moins peu conventionnel. Ensuite pour avoir su me guider et rediriger les efforts quand il a fallu le faire. Et enfin pour leur conseils et présence tout au long de ces trois ans. Je veux aussi remercier tous ceux qui ’ot apporté ue aide préieuse sur le pla sietifiue ou tehiue. Mike Daso ’a aidé à foruler et orgaiser e ui était ou des éduses à zooathelles e ue fore puliale à lauelle j’aurais eu ie du al à arrier seul. Je remercie le gouereet de Palaos de ’aoir peris d’éhatilloer das les las aris de l’arhipels et les membres du CRRF pour leur aide logistique. Je remercie Oanez Lebeau et Rudolph Coraisier, pour ’aoir foré à l’isotopie stale ; Antoine Bideau et Fabienne Le Grad pour ’aoir foré à l’étude des aides gras. Grâe à ous, j’ai pu oeer à leer le oile sur la utritio des éduses à zooathelles. Je reerie aussi Eri Daas et l’éuipe d’Oéaopolis pour leur aide dans mes tentatives plus ou moins fructueuses de maintenir des méduses en aquarium afin de les étudier. Je reerie aussi eu ui ’ot aidé sur le pla scientifique mais dont les efforts sont moins visibles dans cette thèse, simplement parce que des pistes ot été eplorées et aadoées, ou ’ot pas aouti. C’est otaet le as de Laure Peuerie pour ’aoir foré à la odélisatio DEB si si, o a teté !). Je pense aussi à Jonathan Flye-Sainte-Marie pour so aide das les sstes d’oétrie et pour les riolages de kreisels. Je reerie aussi es stagiaires, ui ’ot aidé sur ertains points pour ma thèse, ou ont exploré des sujets connexes : Océane, Enora, Thomas, Sandra, Garance et Nolwenn. Je pese u’o ’aura jamais autat etedu parler de éduses à l’uiersité de Brest, et ’est e oe partie grâe à ous ! Je remercie aussi bien sûr tous es ollgues du lao ae ui j’ai pu e lier pedat es trois ans, Kévin, William, Leslie, Chloé, Elyne, Natalia, Alex, Gaëtan, Jean-François, Houda, Pauline, Antoine, Fanny, Fanny, Aurélien, Sarah, Julien, Manon, Philippe, Mariana, Morgan, Lucien et j’e oulie. Un remerciement tout particulier à mes deux collègues de bureau principaux (les autres on les a moins vus !) Justine et Jordan pour leur amitié et leur capacité à supporter III l’eploi du ot « strobilation » à raison de dix fois par jour. Jorda, j’espre u’o aura d’autres oasios d’aller s’assourdir au Hellfest ou ailleurs ! Un grand merci, bien sûr, à ma bande de potes, ils se reconnaitront ! Ils ont toujours été là pour oi, et je pese ue ’est parti pour otiuer oe ça logteps. Meri d’tre ous ! J’e profite pour eoer tout o soutie et o aitié à Lola, elle aussi earuée das une thèse mais cette fois, en Ecosse. Finalement, u grad eri à a faille, si j’e suis là ’est grâe à ous. Depuis l’époue où je ’étais u’u tout petit hiard, ous ’aez tous eouragé à tre urieu et à déourir le ode. Tout ça ’a logiueet poussé ers la siee et e oilà ae u ausrit de thèse entre les mains ! IV Note sur la Traduction Française Ce manuscrit est écrit essentiellement en anglais. Toutefois, certaines sections sont traduites en français : Les résumés des différents chapitres. Les légendes des figures. Les légendes des tableaux. Les titres des sections principales. Un résumé développé en français est également donné dans les pages suivantes, en avant- propos au corps du document de thèse Note on the French Translation This manuscript is written primarily in English. However, parts of it are translated in French: The abstracts of the different chapters. The figure captions. The table captions. The titles of the main sections. A detailed abstract is also given in French in the following pages as a foreword to the manuscript V Résumé Développé 1. Introduction : Rôle des Méduses dans les Ecosystèmes Marins Les méduses (ici employé au sens large de cnidaires pélagiques) ont historiquement été considérées oe ’aat u’u rôle égligeale das les éosstes aris. Cette isio est à préset dépassée. La reherhe des derires dizaies d’aées portant sur ces organismes a en effet démontré leur importance et leurs rôles dans les écosystèmes marins. Coe d’autres prédateurs pélagiues, les méduses sont à présent considérées comme des membres à part entière des réseaux trophiques, pouvant intervenir sur la dynamique des communautés, les les des utriets ou l’eport de matière organique ers l’oéan profond. Toutefois, un certain nombre de caractéristiques distinguent les méduses des autres prédateurs pélagiues. L’une de ces caractéristiques notable est la tendance de certaines espèces de méduses à former des blooms. Un bloom de méduses est défini comme une augetatio iportate de l’effetif de la population de ces organismes en lien avec leur phénologie. Deux caractéristiques fondamentales des méduses expliquent leur capacité à former ces blooms : (1) La foratio d’u loo éessite u fort recrutement de jeunes méduses. Cela est assuré, chez la plupart des espèces, par la transition de la phase polype à la phase méduse (processus de strobilation chez les scyphozoaires). (2) Ces jeunes recrues doivent ensuite grandir rapidement pour que la populatio atteige d’iportates ioasses. Chez les éduses, ette roissae rapide est perise par leur pla d’orgaisatio simple et leur importante teneur en eau (> 95 %). Tous es aspets de l’éologie et de la daiue des populatios de éduses sot toutefois assujettis à leur nutrition ; i.e. à la capacité des individus à trouver dans leur environnement les ressources nécessaires à leur croissance et leur reproduction. La plupart des méduses sont striteet prédatries, s’alietat généralement sur du micro- ou du mésozooplancton. Aisi, la foratio d’u loo de éduses est souet orrélée à u pi das la uatité de proies dispoiles. Toutefois, de oreuses espes de éduses disposet d’ue seode source de nutrition via une photosymbiose avec des dinoflagellés autotrophes (zooxanthelles). Ce type de symbiose est généralement considéré comme mutualiste. Les méduses fournissent aux zooxanthelles un abri et un accès privilégié à leurs produits VI d’erétio. Les zooathelles fourisset au éduses une part importante des produits de la photosynthèse. La combinaison de (1) la nutrition hétérotrophe des méduses (prédation de zooplancton), de (2) la nutrition autotrophe des zooxanthelles (photosynthèse), et 3 d’u échange de nutriments entre les deux partenaires, implique que ces méduses à zooxanthelles peuet tre osidérées oe iotrophes e tat u’holoiotes (i.e. hôte et symbiontes considérés comme un seul organisme). L’étude de l’éologie des méduses à zooxanthelles et de leur nutrition mixotrophe constituent les thématiques principales de cette thèse organisée en quatre chapitres résumés ci-après. 2. Chapitre I : Revue de la Diversité, des Traits de Vie, et de l’Ecologie des Méduses à Zooxanthelles Le premier Chapitre de cette thèse présente une revue de la littérature concernant la diersité, les traits de ie, et l’éologie des méduses à zooxanthelles. 2.1. Diversité des Méduses à Zooxanthelles et de leurs Symbiontes Au moins sept apparitions de la symbiose médusozoaire-zooxanthelles peuvent être identifiées au ours de l’histoire éolutie des édusozoaires. Deux concernent des groupes d’hdrozoaires e présetat pas de phase éduse taos Filifera I & II, et Marooloia. Les cinq autres apparitions concernent des taxons contenant des méduses, et se répartissent dans tous les principaux groupes de édusozoaires à l’eeptio des staurozoaires : Une apparition est recensée chez les cubozoaires (taxon Carybdeida), deux apparitions chez les hydrozoaires (taxons Capitata et Laodiceida), et deux apparitions chez les scyphozoaires (taxons Coronatae et Kolpophorae). Dans la plupart des cas, les méduses à zooxanthelles sont apparemment peu diversifiées. Une exception importante toutefois est le taxon des Kolpophorae, qui est diversifié (ca. 40-50 espèces) et pourrait ne contenir que des espèces à zooxanthelles. Cela implique que 20 à 25 % des scyphozoaires sont des espèces à zooxanthelles (symbiontes facultatifs inclus). Chez les scyphozoaires, la nutrition mixotrophe ’est do pas un caractère exceptionnel mais est, au contraire, plutôt commune. VII Les zooxanthelles présentes dans les méduses à zooxanthelles sont généralement des Symbiodiniaceae (principalement les genres Symbiodinium et Cladocopium). Plus rarement, d’autres taos peuet tre troués, otaet des zooathelles de la faille des Thoracosphaeraceae. 2.2. Traits de Vie Communs aux Méduses à Zooxanthelles Les méduses à zooxanthelles, bien que diverses phylogénétiquement, semblent partager certaines caractéristiques communes.
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