Animal-Mediated Seed Dispersal and Tropical Rainforest Regeneration in the Face of Anthropogenic Activities in a Forest- Savanna

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Animal-Mediated Seed Dispersal and Tropical Rainforest Regeneration in the Face of Anthropogenic Activities in a Forest- Savanna Dissertation présentée en vue de l’obtention du grade de Docteur en Sciences Année académique 2016-2017 Franck Trolliet Dispersion des graines zoochores et régénération de la forêt tropicale humide face aux activités anthropiques dans une mosaïque de forêts-savanes en République Démocratique du Congo ••• Animal-mediated seed dispersal and tropical rainforest regeneration in the face of anthropogenic activities in a forest- savanna mosaic in Democratic Republic of the Congo Université de Liège Département de Biologie, Ecologie, Evolution Unité de Biologie du Comportement Disseration présentée en vue de l’obtention du grade de Docteur en Sciences, spécialité Biologie des Organismes et Ecologie Année académique 2016-2017 Franck Trolliet Dispersion des graines zoochores et régénération de la forêt tropicale humide face aux activités anthropiques dans une mosaïque de forêts-savanes en République Démocratique du Congo Sous la supervision d’Alain Hambuckers JURY Prof. Jean-Pierre Thomé, Université de Liège Président Prof. Jean-Louis Doucet, Gembloux Agro-Bio Tech / Université de Liège Secrétaire Dr. Marie-Claude Huynen, Université de Liège Rapporteur Prof. Pierre-Michel Forget, Muséum d’Histoire Naturelle de Paris Rapporteur Dr. Eike Neuschulz, Biodiversity and Climate Research Centre (BiK-F) Rapporteur A ma famille, Jantje et Jules 6 Remerciements Remerciements Je souhaite tout d’abord adresser mes remerciements aux personnes sans qui la présente thèse n’aurait pu voir le jour. C’est initialement grâce à Marie-Claude Huynen que j’ai pu entendre parler de ce projet de recherche passionnant en Afrique Centrale. Je souhaite te remercier pour ta présence, tes encouragements et la confiance que tu m’as accordée depuis mes premiers moments à Liège. C’est Alain Hambuckers qui m’a ensuite permis de prendre ce projet en main. Je me souviendrai longtemps de mon enthousiasme face à l’idée de mettre en place un projet sur la régénération forestière dans la brousse congolaise. J’ai rapidement dû m’adapter face à la multitude de possibilités qui m’étaient offertes et de challenges qui me barraient la route. Dès le début donc, ce projet a été énormément formateur et enrichissant. Alain, tu t’es montré incroyablement disponible pour écouter mes doutes, discuter, et me donner des conseils avisés pour avancer, étape par étape, tout en m’accordant beaucoup de confiance et de libertés quant aux choix que je faisais. Je t’en remercie grandement! Je remercie grandement la Politique Scientifique Fédérale Belge (Belspo) d’avoir financé ce projet pendant les deux premières années, le F.R.S.-FNRS pour m’avoir attribué une bourse FRIA les trois années suivantes, ainsi que le Patrimoine de l’ULg pour m’avoir appuyé financièrement pour que je puisse mener mes missions sur le terrain et participer aux congrès scientifiques. Je tiens à remercier les membres de mon comité de thèse, Marie-Claude Huynen, Roseline Beudels-Jamar, Louis François, et Pierre-Michel Forget, pour leur présence régulière et leur patience lors de mes présentations quelques peu redondantes, ainsi qu’à leurs nombreux conseils et encouragements. Pierre-Michel, avant notre rencontre, j’ai eu la chance de choisir une espèce de Myristicaceae pour mon projet, le lien était fait! Tu as rejoint le bateau en cours de route mais tu as été une personne clé pour ma thèse, et ton implication est loin de s’être limitée aux réunions annuelles du comité. Tu as toujours été partant pour prendre une journée pour venir à Liège et discuter de mes résultats plus ou moins satisfaisants malgré ton emploie du temps très chargé. Tes connaissances pointues ont sans aucun doute largement contribué à améliorer la qualité des différents chapitres. Tu m’as aussi poussé à donner une suite ambitieuse à la deuxième partie du projet. Merci! J’espère de tout cœur que cela portera ses fruits et que notre collaboration ne fait que commencer. Je voudrais remercier toutes les personnes qui m’ont accompagné pendant les différentes missions au Congo. Adeline Serckx, africaine blanche parfaitement organisée qui m’a appris dès mes premiers jours sur le continent à prévoir aussi bien des câpres et du parmesan pour agrémenter nos repas en brousse, que de gérer une équipe de pisteurs locaux pour récolter les données vitesse grand V. Jief, que dire? Passer de délires les plus absurdes, à des discussions sur la dynamique de succession forestière, et à l’organisation logistique pour aller camper dans «la savane aux éléphants» fût plutôt riche en émotions. Vous avez tous les deux été des personnes importantes pendant ma thèse, que ce soit pour les moments partagés sur le terrain que pour m’avoir inspiré et motivé dans mes recherches. Merci aussi à Ménard Mbende et Céline Devos pour leur accueil et leur aide précieuse à mes débuts à Malebo. Un tout grand Remerciements 7 merci à Jean Mobuli aka chef de base pour son aide et ses conseils toujours sages et précieux, ainsi que sa chaleureuse présence pendant les longs mois à Guantana-Malebo. Merci à toute l’équipe de la base de Malebo. Chab et Olivia, vous y êtes arrivés seulement à la fin de mon terrain, mais quel plaisir de vous avoir eu à mes côtés pour ma dernière mission! Je tiens aussi à remercier Paulin et Wivine de la ferme Mbou-Mon-Tour, ainsi que les chefs-de-terres des villages de Nkala, Embirima, Mpelu, et Mbanzi pour m’avoir autorisé à mener mes recherches dans leurs forêts. Toutes les données acquises résultent des efforts de nombreux pisteurs que je tiens à remercier chaleureusement : Mora, Pala, Ngamba, Americain, Nioka, Jean-Baptiste, Boula, Papa Denis, Idia, Mpesse. Merci également aux mémorantes Kim Mathy et Alice Dauvrin de m’avoir aidé pour la récolte de données. Je tiens aussi à remercier les équipes du WWF et de Mbou-Mon-Tour à Kinshasa pour leur support logistique. Enfin, merci à la moto Buxer de m’avoir permis des allez retours incessants entre mes nombreux sites d’études. Je voudrais remercier l’équipe des forestiers de Gembloux Agro-Bio Tech. Tout d’abord Cédric Vermeulen pour ton idée judicieuse de publier un article sur les camera-trap. Jean-Louis Doucet, pour ta disponibilité pour discuter de mes différents objectifs et méthodes. Tu as toujours fait des remarques constructives en mettant le doigt sur les points faibles de mon projet. C’est en fin de thèse que j’ai eu la grande chance de bénéficier de tes compétences de botaniste pour identifier mes centaines d’échantillons de plantes. Avec Jean-François Gillet et Olivier Lachenaud, vous avez été une ressource inestimable pour valoriser mes données. Je vous en remercie. Merci aussi aux différents membres de l’équipe de Foresterie Tropicale pour leur accueil lors de mes journées de transit à Gembloux. J’ai durant cette thèse rencontré plusieurs personnes d’une grande richesse qui m’ont toujours inspiré et accompagné, que ce soit en Belgique ou ailleurs sur la planète lors de conférences: Laurence, Fany, Barbara, Nima, Lauranne, Yaëlle. Je tiens aussi à remercier Thomas Drouet et David Bauman pour la collaboration de dernière minute et les analyses des échantillons de sols et les statistiques quelques peu précipités. Mais un tel projet n’est pas simplement le fruit de collaborations scientifiques, de kilomètres en forêt, et de longues heures passées devant Word et Excel. C’est aussi et sans nul doute grâce à un soutien extérieur que j’ai eu la chance de mener ce bout de chemin et de conduire à bien mon projet tout en restant équilibré. Je voudrais ainsi remercier tout particulièrement ma moitié pour son respect dans mes choix et son incroyable patience pendant mes périodes d’absences souvent trop longues et épicées à coup de Plasmodium falciparum, ainsi que mes périodes de confinement devant l’ordinateur parsemées de « j’ai pas le temps je dois bosser ». Jantje, tu m’a toujours accompagné et encouragé pour que je poursuive dans ma voie, et ce malgré les compromis à faire. Pendant ces cinq années, tu m’as aidé à relativiser et à garder les pieds sur terre pendant les périodes difficiles, les départs, les retours, les «après». Et, parlant sans cesse de graines qui germent et de jeune plantes qui poussent, je me devais moi-même de faire cette magnifique expérience biologique et de m’investir ailleurs que dans la quête interminable de résultats significatifs: notre petit Jules en est la plus belle des concrétisations. Merci milles fois pour tout. Je souhaite finalement remercier profondément ma famille pour leurs encouragements et leur soutien tout au long de ce projet. 8 Résumé Résumé Les patrons de biodiversité dans les forêts tropicales humides résultent d’interactions complexes à l’échelle de la communauté, tels que la dispersion des graines par les animaux. Les activités anthropiques (chasse, perte et fragmentation des forêts, etc.) perturbent ces processus écologiques et menacent ainsi le maintien de la diversité à l’échelle de la communauté végétale. Néanmoins, le niveau de résilience des processus impliqués dans la régénération forestière dans les paysages perturbés demeure incertain. La présente thèse vise à comprendre comment les services de dispersion des graines assurés par les grands frugivores ainsi que leur rôle fonctionnel dans la régénération des plantes sont affectés dans un paysage influencé par les activités anthropiques. Nous avons mené un travail de terrain dans une mosaïque de forêts- savanes en République Démocratique du Congo, un paysage ayant une structure spatiale fragmentée et qui abrite des activiés anthropiques (chasse, agriculture itinérante sur brûlis). Premièrement, nous avons examiné les services de dispersion des graines assurés par un grand primate menacé, le bonobo (Pan paniscus). Les bonobos dispersent une majorité de grandes graines que peu d’autres frugivores peuvent disperser, qui incluent de nombreuses espèces sciaphiles.
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