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Ecology, Forms and Functions of the Basal Angiosperms from New Caledonia Santiago Trueba

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Ecology, Forms and Functions of the Basal Angiosperms from New Caledonia Santiago Trueba Ecology, forms and functions of the basal angiosperms from New Caledonia Santiago Trueba To cite this version: Santiago Trueba. Ecology, forms and functions of the basal angiosperms from New Caledonia. Life Sciences [q-bio]. Universit´ede Montpellier, 2016. English. <tel-01392906> HAL Id: tel-01392906 https://tel.archives-ouvertes.fr/tel-01392906 Submitted on 4 Nov 2016 HAL is a multi-disciplinary open access L'archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destin´eeau d´ep^otet `ala diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publi´esou non, lished or not. The documents may come from ´emanant des ´etablissements d'enseignement et de teaching and research institutions in France or recherche fran¸caisou ´etrangers,des laboratoires abroad, or from public or private research centers. publics ou priv´es. Délivré par l' UNIVERSITÉ DE MONTPELLIER Préparée au sein de l’école doctorale GAIA Et de l’unité de recherche AMAP Spécialité : Écologie, É volution, Ressources Génétiques, Paléobiologie Présentée par Santiago Trueba -Sánchez Ecology, forms and functions of the basal angiosperms from New Caledonia Ecologie formes et fonctions des angiospermes basales en Nouvelle-Calédonie Soutenue le 26 avril 2016 devant le jury composé de : Daniel BARTH ÉLÉMY, DR, CIRAD Directeur de thèse Tim J. BRODRIBB, A ssociate Professor , University of Tasmania Rapporteur Hervé COCHARD, DR, INRA Examinateur Pierre -Eric LAURI, Ingénieur hors classe , INRA Examinateur Porter P. LOWRY II, Senior Curator , Missouri Botanical Garden Rapporteur Doyle McKEY, Professeur, Université de Montpellier Examinateur Ecology, forms and functions of the basal angiosperms from New Caledonia Santiago Trueba -Sánchez Director : Daniel Barthélémy (CIRAD , France) Advisors : Sandrine Isnard (IRD , France ) Mark E. Olson (UNAM , Mexico) Jiro Taniguchi, The Walking Man (1995) Que beleza é sentir a natureza Ter certeza pr'onde vai E de onde vem Que beleza é vir da pureza E sem medo distinguir O mal e o bem Tim Maia, Imunização Racional (1974) A mis padres Résumé L’une des remarquables originalités de la flore néo-calédonienne repose sur la présence de nombreuses lignées correspondant aux premières divergences des plantes à fleurs, compte tenu de leurs positions phylogénétiques. Au sein de ces lignées, certaines espèces sont susceptibles de porter des traits morpho- anatomiques ancestraux. Par conséquent, dans un contexte comparatif, l'étude de ces espèces peut fournir des informations cruciales pour comprendre les premières étapes évolutives des angiospermes. Un premier volet de cette thèse vise à étudier des caractéristiques structurelles et fonctionnelles des groupes représentatifs d’angiospermes basales. L'étude d' Amborella trichopoda , espèce sœur de toutes les angiospermes, nous a montré une covariation des traits fonctionnels (tige-feuille) et une plasticité morphologique en réponse à des variations de l'environnement lumineux. Cela suggère que ces réponses plastiques étaient déjà présentes chez l'ancêtre commun de toutes les plantes à fleurs. En parallèle, l’étude de l'évolution anatomique des Piperales, ordre le plus riche parmi les angiospermes basales, suggère que leur ancêtre commun aurait possédé un cambium actif. Ces résultats supportent que les premières angiospermes avaient une forme de vie ligneuse et que la structure sympodiale à été acquise dans les premières étapes évolutives des angiospermes. Les angiospermes basales comptent parmi les lignées d'angiospermes surreprésentées en Nouvelle- Calédonie. Cependant, les mécanismes à l’origine de cette dysharmonie demeurent inexplorés. Un deuxième volet de cette thèse analyse la répartition environnementale des angiospermes basales de l'archipel afin de connaitre leurs préférences en termes d’habitats, ainsi que leurs exigences environnementales. Cette distribution environnementale a également été analysée au regard de leur résistance à la sécheresse. Nous montrons que la plupart des espèces présentent une préférence marquée pour des habitats de forêt humide avec des faibles variations en température. La vulnérabilité hydraulique face à la sécheresse apparait comme un trait majeur qui confine la distribution de ces espèces dans des habitats humides. Ces conditions auraient persisté dans des zones refuges dans l'archipel lors de la dernière période glaciaire, permettant ainsi le maintien de certaines lignées d’angiospermes basales. Une stabilité climatique passée pourrait donc être à l'origine de la surreprésentation de certains groupes d'espèces forestières qui ont disparu dans les régions voisines. La distribution des angiospermes basales néo-calédoniennes, ainsi que leur sensibilité à la sécheresse, supportent l'hypothèse suggérant que les premières angiospermes habitaient des milieux humides et stables. Mots-clés: anatomie du bois, angiospermes basales, biogéographie, écologie fonctionnelle, écophysiologie, évolution, Nouvelle-Calédonie, vulnérabilité à la sécheresse. Abstract One of the remarkable characteristics of the New Caledonian flora is the presence of numerous angiosperm lineages recognized as the earliest divergences of the flowering plants, due to their phylogenetic positions. Within these lineages, some species are likely to bear ancestral morpho-anatomical features. Therefore, under a comparative perspective, the study of these species can provide compelling information for understanding the early evolutionary stages of angiosperms. The first part of this thesis aims to study the structural and functional characteristics of representative groups of basal angiosperms. The study of Amborella trichopoda , sister species to the remaining flowering plants, shows a covariation of functional traits (stem-leaf) and a morphological plasticity in response to changes in the light environment. This suggests that these plastic responses were already present in the common ancestor of angiosperms. In parallel, the study of the anatomical evolution of Piperales, the most diversified basal angiosperm order, suggests that their common ancestor had an active cambium. These results support the hypothesis that early angiosperms had a woody habit and that sympodial growth may have been acquired early during angiosperms evolution. Basal angiosperms are among the over-represented angiosperm lineages of New Caledonia. However, the mechanisms underlying this disharmony remain unexplored. A second component of this thesis analyzes the environmental distribution of New Caledonian basal angiosperms to know their habitat preferences and their environmental requirements. Further, we assess the influence of their drought tolerance on their environmental distribution. We show that most species have a preference for rain forest habitats with small variations in temperature. Drought-induced hydraulic vulnerability stands as a major trait that restricts the distribution of these species to humid habitats. These stable conditions seem to have persisted in refugial areas in the archipelago during the last glacial maximum, allowing the persistence of basal angiosperm species. Therefore, a past climatic stability could explain the over-representation of some groups of forest species that may have disappeared in neighboring regions. The distribution of basal angiosperms in New Caledonia, as well as their drought sensitivity, support the hypothesis suggesting that early angiosperms lived in humid and stable environments. Keywords: basal angiosperms, biogeography, drought vulnerability, ecophysiology, evolution, functional ecology, New Caledonia, wood anatomy. REMERCIEMENTS Tout d’abord, je tiens à remercier mes encadrants de m’avoir fait confiance tout au long de ce projet. Merci à Sandrine Isnard de m’avoir donné l’opportunité de mener un projet de thèse passionnant et de m’avoir ainsi permis de découvrir la richesse de la flore de Nouvelle-Calédonie. Merci à Mark Olson pour nos discussions sur l’évolution des plantes et pour ses nombreux conseils de rédaction. Je remercie également Daniel Barthélémy pour nos échanges sur l’architecture d’ Amborella et pour la supervision de mes travaux. Je remercie les rapporteurs, Tim Brodribb et Pete Lowry, ainsi que les membres examinateurs, Hervé Cochard, Pierre-Eric Lauri et Doyle McKey pour leur intérêt et pour l’honneur qu’ils me font de participer à mon jury de thèse. Je tiens à remercier particulièrement Robin Pouteau sans qui le chemin n’aurait pas été le même. Merci pour ton soutien à chaque étape de cette thèse, pour nos discussions sur le projet, mais surtout merci pour ton amitié. M āuruuru Robin! Un énorme merci à mes collègues du labo de botanique, qui m’ont soutenu les trois dernières années. Je remercie Thomas Ibanez pour son soutien dans la révision de ce manuscrit de thèse, merci aussi Tom pour tous les bons moments passés ensemble sur le caillou. Un grand merci à Laure Barrabé pour son inestimable soutien et ses nombreuses relectures, merci aussi de m’avoir changé les idées avec tes blagues journalières. Je remercie mon amie Elodie Blanchard d’être toujours à l’écoute, merci Elo pour tes conseils pertinents et d’avoir partagé avec moi la vie de thésard. Un grand merci aussi à Philippe Birnbaum pour son soutien, son énergie débordante, et sa motivation contagieuse. Merci à Vanessa Hequet de toujours veiller à ce que tout se passe bien pour moi et pour son rire qui résonne dans le couloir. Merci à Dimitri Justeau pour nos discussions à la lumière tamisée du Nakamal. Merci à
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