Dispersion Épizoochore Des Graines Par Les Ongulés Sauvages Dans Des Paysages Changeants : Le Cerf Élaphe Comme Étude De Cas Ushma Shukla

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Dispersion Épizoochore Des Graines Par Les Ongulés Sauvages Dans Des Paysages Changeants : Le Cerf Élaphe Comme Étude De Cas Ushma Shukla Dispersion épizoochore des graines par les ongulés sauvages dans des paysages changeants : le cerf élaphe comme étude de cas Ushma Shukla To cite this version: Ushma Shukla. Dispersion épizoochore des graines par les ongulés sauvages dans des paysages changeants : le cerf élaphe comme étude de cas. Sciences agricoles. Université d’Orléans, 2020. Français. NNT : 2020ORLE3060. tel-03166043 HAL Id: tel-03166043 https://tel.archives-ouvertes.fr/tel-03166043 Submitted on 11 Mar 2021 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. UNIVERSITÉ D’ORLÉANS ÉCOLE DOCTORALE Santé, Sciences Biologiques et Chimie du Vivant INRAE Val de Loire, UR Écosystèmes Forestiers THÈSE présentée par Ushma SHUKLA soutenue le : 24 Septembre 2020 pour obtenir le grade de : Docteur de l’Université d’Orléans Discipline/ Spécialité : Écologie Epizoochorous seed dispersal by wild ungulates in changing landscapes the red deer as a case study CO-DIRECTEURS BALTZINGER Christophe IDAE, HDR, INRAE Val de Loire, Université d’Orléans REINEKING Björn Directeur de recherche, INRAE LESSEM, Université Grenoble-Alpes RAPPORTEURS LOISON Anne Directeur de recherche, CNRS, LECA, Université Savoie Mont Blanc DUPOUEY Jean-Luc Directeur de recherche, INRAE, UMR Silva, Université de Lorraine JURY DUPOUEY Jean-Luc Directeur de recherche, INRAE, UMR Silva, Université de Lorraine (Président du jury) LOISON Anne Directeur de recherche, CNRS, Université Savoie Mont Blanc BANKHEAD Stephanie Maître de conférences, Université d’Orléans RIOTTE-LAMBERT Louise Post-doctorante, Université de Glasgow PAYS-VOLARD Olivier Maître de conférences, HDR, Université d’Angers BALTZINGER Christophe IDAE, HDR, INRAE Val de Loire, Université d’Orléans REINEKING Björn Directeur de recherche, INRAE LESSEM, Université Grenoble-Alpes Epizoochorous seed dispersal by wild ungulates in changing landscapes : the red deer as a case study Summary Epizoochory is one mode of plant dispersal, where seeds are conveyed externally on animals. It mostly concerns forbs and graminoids, shrubs and trees relying more on endozoochory or abiotic vectors. Epizoochory is considered a strong biotic filter that shapes local plant assemblages from the regional pool of species. Abundant populations of large-bodied ungulates and their home range movements provide frequent ‘mobile links’ among habitat patches. For my thesis, I was interested in assessing the role of wild ungulates, especially red deer, in epizoochorous seed dispersal. I addressed 1) the role of ungulates in structuring local plant communities with respect to dispersal-related plant traits, 2) the effect of landscape composition on movement decisions made by individual red deer on subsequent seed dispersal kernels, and 3) the complementarity of ungulate-mediated dispersal modes and their overall influence on local plant communities. I used a combination of empirical field data and ecological modeling to address these questions. I found that zoochory presents a weak signal in structuring plant communities compared to other abiotic factors, but that dispersal-enabling traits were good predictors. I demonstrated the importance of accounting for home ranging behavior in range-resident ungulates in the seed dispersal kernel, which prevents from over-estimating dispersal distances. Moreover, I underlined how considering the total seed load dispersed by individual red deer is crucial and reveals the complementarity of dispersal modes in the total dispersal kernel. Finally, I highlighted the underrated role of ungulates in spreading invasives as a real threat to biodiversity. Keywords: seed dispersal kernel, functional traits, complementarity, changing landscapes, plants-ungulate interactions Dispersion épizoochore des graines par des ongulés sauvages dans des paysages changeants : le cerf élaphe comme étude de cas Résumé L'épizoochorie définit la dispersion des plantes véhiculées sur le corps des animaux. Elle concerne herbacées et graminées, arbres et arbustes étant dispersés par endozoochorie et vecteurs abiotiques. Filtre biotique sélectif, elle façonne les communautés végétales locales à partir du pool régional d'espèces. Les populations abondantes d’ongulés sauvages et leurs longs déplacements créent ainsi de fréquents "liens mobiles" entre patchs d'habitat. J’ai ainsi étudié le rôle de ces animaux, notamment du cerf élaphe, comme vecteurs de dispersion épizoochore dans la structuration des communautés végétales locales en tenant compte des traits des plantes favorisant la dispersion. Puis, j’ai évalué comment structure et composition du paysage affectaient les déplacements individuels du cerf, les distances de dispersion et le site de dépôt des graines. Finalement, j’ai comparé différents modes de dispersion à l’échelle du vecteur individuel. Pour répondre à ces questions, j'ai combiné données empiriques de terrain et approches de modélisation écologique. J'ai constaté que la zoochorie constitue un signal faible, par rapport aux facteurs abiotiques, dans la structuration des communautés végétales, mais les traits facilitant la dispersion sont de bons prédicteurs. J'ai montré l'importance de tenir compte du domaine vital individuel pour ne pas surestimer les distances de dispersion. En outre, j'ai démontré qu’il fallait considérer la charge totale en graines véhiculées car elle révèle la complémentarité des modes de dispersion. Pour finir, j'alerte sur le rôle sous-estimé des ongulés sauvages et domestiques dans la propagation des plantes exotiques, réelle menace pour la biodiversité Mots clés : dispersion des graines, traits fonctionnelles, complémentarité, paysages changeants, interactions plantes-ongulés INRAE Centre Val de Loire Domaine des Barres Nogent-sur-Vernisson 45290 France ACKNOWLEDGEMENTS There is an old saying “It takes a village to raise a child”. So, I would like to thank the people who helped me raise this thesis baby :) Project COSTAUD and Irstea (now, INRAE) for funding this PhD and giving me the opportunity to follow my research interests freely. My supervisors, Christophe Baltzinger and Björn Reineking. Your unwavering patience and support helped me to sail through big, bumpy waves all the way to the finish line. My sincerest gratitude to both of you for mentoring me. I would like to thank the members of my academic committee- Audrey Muratet, Aurélie Coulon and Carsten Eichberg for lending a patient ear to my research ideas and for helping to smooth out the creases. I would like to thank all the members of the jury who agreed to evaluate and review the thesis. I would like to thank the department and staff at the Hunting and Forest office of Domaine National de Chambord for the permits to work there and support during field work. I would also like to thank my collaborators, Sonia Said and Marco Heurich as well as fellow PhD students Agathe Chassagneux and Maik Henrich. Thanks are due to Frédéric Archaux, Director of the Nogent centre, for the innumerable little ways in which he always helped me. My colleagues at INRAE-Nogent. Especially the highly commendable job that the general services team does. Sylvie Le Roux, Florance Van Den Boom and Jean-Pascal Barrau always helped with tricky paperwork. Richard Chevalier, for helping out with field and greenhouse work and teaching me what good data management and archiving looks like. Hai Linh Nguyen for all her hard-work during her internship. My team mates in FONA. Anders Mårell for all the sage advice and support. Mathilde Brodut and Adélie Chevalier for being the strong pillars on which our research endeavors depend and always rising to the occasion. Agnès Rocquencourt for being my go-to person for all kinds of mundane things. Jean-Pierre Hamard and Yves Boscardin, for their support. Nadège Bonnot and Laura Chevaux for their friendship and the endless rounds of coffee-break ;) Phillippe Guillemard and Hilaire Martin, colleagues-turned-friends who were there, at the drop of a hat, be it solving a technical problem with some software or just needing a coffee break or giving me life lessons. Fabien Laroche and Yoan Paillet for always fielding my silly questions for all things quantitative with immense patience. Yoan, also, for hosting my stay in Grenoble. Guilhem Parmain, for all the pampering :) Marie Baltzinger with whom I could strike up a conversation about nearly everything. Cécile Joyeau for always coming up with new plans to break away from the routine. Jordan Bello for all the laughs. Tiffani and Ugoline, the two bright rays of sunshine that can brighten up any dull day. I may not be able to find the right words to express myself, but I am grateful for your friendship. Friends in India and in the world at large, who don’t let geography get in the way of our friendship. Shraddha, Harsha, Apeksha, Darshan, Mrunal, Akanksha, Geetha, Karpa, Rajat, Ipsita, Dharma, Rutuja, Amit, Neelamda, Rohit, Divya, Chethi, Rafael and Camille. Thank you for the care, concern and all the good times. To AK and NMN, the two teachers who are always there to guide my path. My parents who always encouraged me to follow wherever
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