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Università degli Studi di Ferrara

DOTTORATO DI RICERCA IN

BIOLOGIA EVOLUZIONISTICA E AMBIENTALE

co-tutela di tesi con Université de Montpellier

CICLO 27

COORDINATORE Prof. Guido Barbujani

Scale-dependent approaches in

conservation biogeography of a cosmopolitan raptor: the Osprey.

Settore Scientifico Disciplinare BIO/05

Dottorando Tutore Universitá di Ferrara Dott.

Monti Flavio Prof. Fusani Leonida

(firma) (firma)

Tutore Université de Montpellier

Prof. Duriez Olivier

(firma) A n n i 2 0 1 2 2 0 1 4 2 3

Délivré par Université de Montpellier

Préparée au sein de l"école doctorale SIBAGHE-GAIA

Et de l"unité de recherche UMR5175 CEFE

Présentée par Flavio Monti

Soutenue le 10/04/2015 devant le jury composé de Dr. Olivier Duriez, Maitre de Conf., Univ. de Montpellier Directeur de these Pr. Leonida Fusani, Professeur, Univ. de Ferrara Directeur de these Pr. Franois Sarrazin, Professeur, Univ. Pierre et Marie Curie Rapporteur Pr. Matteo Griggio, Maitre de Conf., Univ. de Padova Rapporteur

Scale-dependent approaches in

conservation biogeography of a cosmopolitan raptor: the Osprey. 4 5

Flavio Monti

Scale-dependent approaches in

conservation biogeography of a cosmopolitan raptor: the Osprey.

PhD Thesis

Defended on 10 April 2015, University of Ferrara

Jury committee:

Pr. Leonida Fusani - Supervisor - University of Ferrara - Italy Dr. Olivier Duriez - Supervisor - Université de Montpellier - France Pr. Franois Sarrazin - Reviewer - Université Pierre et Marie Curie, Paris - France Dr. Matteo Griggio- Reviewer - University of Padova - Italy

Doctor Europaeus

Pr. Anders P. Tøttrup - Reviewer - University of Copenaghen - Denmark 6

This PhD project was carried out in the Department of Life Sciences and Biotechnology of the University of

Ferrara (Italy), and in the Department of Biogeography and Ecology of Vertebrates, Centre d'Ecologie

Fonctionnelle et Evolutive (CNRS UMR5175), University of Montpellier (France), according to the joint

collaboration signed by the two institutions.

The production of this thesis was financially supported by the Foundation Prince Albert II de Monaco and the

Associazone Italiana della Fondation Prince Albert II de Monaco ONLUS, the Corsica Natural Regional Park

(France), the Maremma Regional Park Agency (Italy), the Mediterranean Small Islands Initiative PIM

coordinated by the Conservatoire du Littoral (France), the Haut Commissariat aux Eaux et Forêt et à la Lutte

Contre la Désertification and Association de Gestion Intégrée des Ressources (AGIR) (Morocco). FM benefits

from a grant from the Maremma Regional Park and from a mobility grant from the Universitá Italo

Francese/Université Franco Italienne.

Collaborations:

David Grémillet -

Centre d"Ecologie Fonctionnelle et Evolutive (CNRS UMR5175)

Claudine Montgelard -

Centre d"Ecologie Fonctionnelle et Evolutive (CNRS UMR5175)

Andrea Sforzi -

Maremma Natural History Museum, Grosseto, Italy

Jean-Marie Dominici - Rèserve Naturelle Scandola, Parc Naturel Règional de Corse, Galeria, France

Thomas Alerstam - Migration Ecology Group, Department of Animal Ecology -Lund University, Sweden

Raymond H.G. Klaassen -

Animal Ecology Group, Centre for Ecological and Evolutionary Studies,

University of Groningen, Netherlands

Mathieu Thévenet - Conservatoire du Littoral, Aix-en-Provence, France

Giampiero Sammuri -

Federparchi - Roma, Italy

Rafel Triay Bagur -

IME (Institut Menorquí d"Estudis), Balearic Islands, Spain

Antoni Muñoz Navarro -

Grup Balear d'Ornitologia i Defensa de la Naturalesa (GOB), Balearic Islands, Spain Ilham Bentaleb - Institut des Sciences de l'Evolution de Montpellier (UMR5554), France Gaël Guillou - Université de la Rochelle, France Rémi Choquet - Centre d"Ecologie Fonctionnelle et Evolutive (CNRS UMR5175). Nibani Houssine - Association de Gestion Intégrée des Ressources (AGIR), Al Hoceima, Morocco.

Hamid Rguibi Idrissi -

University Chouaib Doukkali, Faculty of Sciences, El Jadida, Morocco.

Véronique Arnal -

Centre d"Ecologie Fonctionnelle et Evolutive (CNRS UMR5175)

Pierre-Christian Beaubrun -

Centre d"Ecologie Fonctionnelle et Evolutive (CNRS UMR5175) Alo s Robert - École Pratique des Hautes Études (EPHE), Paris, France

How to cite this thesis:

Monti, F. (2015). Scale-dependent approaches in conservation biogeography of a cosmopolitan raptor: the Osprey. PhD thesis. University of Ferrara / University of

Montpellier.

7

Acknowledgments:

I spent most of my life on the shores of the Mediterranean Sea: cradle of many civilizations, the Mediterranean region is an interweaving of cultures, colours, traditions that leave an important mark on those who, like me, had the good fortune to sail its sea, know the voices of its ports, and smell the scent of its islands. I owe to thank the Mare Nostrum and the

people I met along its shores, during this pilgrimage which had the taste of a personal

Odyssey.

It is difficult to accomplish a challenging mission without good “traveling companions": my fortune was to share this experience with an enthusiastic group of people that, in an atmosphere of friendship and collaboration, constantly supported me along the “PhD- journey". I would say thanks to my supervisors Olivier Duriez, Leonida Fusani, David Grémillet and Andrea Sforzi which have equally contributed to lay the foundation for such successful project with their important scientific contribution and practical suggestions. Thank you for untangling the task concerning the joint supervision and for giving me confidence, freedom in work and have passionately supported the ideas of the project since the beginning. Each story has a starting point: how not remember the meeting with David during one of the hottest days in the Corsican summer of 2010, the preliminary brainstorming on ospreys with Olivier in the magnificent scenery of the canyon of the Grand Causses staring at his vultures, the long walks with Leonida on the snowy ancient walls of Ferrara in winter and the talks with Andrea along the banks of the Ombrone river turning the gaze and thoughts, towards the river mouth, the sea... and more far away. Thanks, you had a strong influence on this thesis and you deeply contributed to make such a wonderful experience. Thanks to the three reviewers that will spend time in reading this manuscript: Fran ois Sarrazin, Matteo Griggio and Anders P. Tøttrup. Thanks to the coordinator of the doctoral school of the University of Ferrara, Guido Barbujani and to the administrative personnel, Lorenzo Ravaglia and Lena Fabbri. Massimiliano Lunedei, of the Department of Life Sciences and Biotechnology, was of great help for administrative paperwork. Thanks to the ancient and new director of the Ecole Doctorale Sibaghe (Montpellier), Bernard Godelle and Michel Lebrun and its administrative personnel, Joelle Lopez and Cendrine Jay-Allemand. I also deeply thank the whole organigram of the Fundation Prince Albert II de Monaco for

funding this project and in particular Hélène El Missouri, Project Administrator, for her

involvement. I am also grateful to the whole personnel of the Universitá Italo 8 Francese/Université Franco Italienne; the institution contributed with a mobility grant from the “Bando Vinci". Dr. Clarisse Bouillot followed the administrative procedures. I am very grateful to Thomas Alerstam and Raymond H.G. Klaassen for the inspiring discussions about migration during my stay at the University of Lund (Sweden). Thanks to your previous work on ospreys and all your data you kindly shared, it was possible to compare migratory strategies of different populations and discuss them within an evolutionary context. For me, it was the most fascinating and stimulating chapter of the thesis. Thanks to Michael U. Kemp, of the University of Amsterdam, for tips on his own R package for calculating tailwind. Knud Andreas Jønsson, Anders P. Tøttrup, Kasper Thorup and Ran Nathan for organizing and accepting me in the Dispersal Symposium, during the International Ornithological Conference in Tokyo (Japan) in 2014. They also organized a very nice dinner at a Japanese restaurant. During international meetings and summer schools an Italian colleague and friend, Giacomo Dell"Omo, was always present being of great help for general advices before oral presentations. Special Acknowledgements (samples and ringing network): We thank all the colleagues that kindly provided osprey samples: Pertti Saurola, Harri Koskinen and Juhani Koivu (Finland), Urmas Sellis (Estonia), Aigars Kalv ns (Latvia), Miroslav V. Babushkin and Vladimir V. Ivanovsky (Russia), Martin Haase and Daniel

Schmidt (Germany), Rolf Wahl, Fran

ois Gossmann and Eric Pasquet (France), Luis Palma and Ursula Hofle Hansen (Portugal), Manuel Siverio (Canary islands), Pedro López-Suárez (Cape Verde), Houssine Nibani (Morocco), Rafel Triay (Balearic islands). A special thanks to Yula Kapetanakos and Alan Poole (Cornell lab of Ornithology, Ithaca, USA) for collecting and performing free analyses of 10 samples from USA. Many thanks are also addressed to the following museums and birds" collection curators that allowed us to collect samples from their collection: Mark Adams (Natural History Museum of London; UK), Steven van der Mije and Bechy Desjardins (Netherlands Centre for Biodiversity Naturalis of Leiden), Isao Nishiumi (National Museum of Nature and Science of Amakubo; Japan), Mikhail Kalyakin, Pavel Tomkovitch, Yaroslav Red"kin (Zoological Museum of Moscow; Russia), Guillermo Delgado (Museo de la Naturaleza y el Hombre of Tenerife Island; Spain) and Enrico Borgo (Museo Civico di Storia Naturale Giacomo Doria di Genova; Italy). We also thank the Portuguese Navy and in particular the contra-almirante José Luís Branco Seabra de Melo (Head of Cabinet of the Navy Chief of Staff) and the capitão-de-mar-e-guerra José Jaime Gonçalves Ribeiro (Director of the Vasco da Gama Aquarium) for providing historical 9 samples of Portuguese ospreys. All extractions from museum specimens were performed using the facilities of the platform “ADN dégradé" (Labex CeMEB, Montpellier, France) coordinated by Christelle Tougard. PCR products were sequenced thanks to the “Bibliothèque du Vivant" sequencing project (CNRS-INRA-MNHN). During the “PhD-journey" I made stops in several places. • Corsica = Pace e salute a o populo Corso...Ci campemu!! Jean-Marie Dominici a été un guide de terrain fondamental durant l"ensemble du projet: en homme avisé de la mer et des montagnes, il m"a appris à trouver le bon chemin jusqu"aux nids de balbuzards sur les falaises rhyolitiques, et m"a montré la splendeur naturelle de l"Ile de Beauté. J"ai eu la chance de profiter de sa connaissance de la vie des balbuzards et de la

gestion des aires protégées de Méditerranée. Aucun des résultats que j"ai pu obtenir lors de ce

projet n"aurait été possible sans son aide et sa persévérance, typiquement corse. Je remercie

également l"ancien et le nouveau président du parc, respectivement Jean-Luc Chiappini et Jacques Costa, ainsi que le directeur Philippe Porruncini. Les gardes de la réserve de Scandola Joseph Albertini (l'Altore o u lione di Corsica), Virgile Le Normand, Julien Tavernier,

Saveriu-Antone Rossi et Fran

ois Paolini, ont été une aide et un support indispensable pendant le travail de terrain en Corse. Des travailleurs professionnels pendant la journée et d"amusants camarades au bar le soir. Alain Camoin a été d"une grande aide pour les suivis des couples reproducteurs de la zone de Cap Corse. Entre 2012 et 2014, j"ai reçu l"aide d"Alo s Robert, Martin Delpuech, Charley Chancelier, Gabrielle Valesi, Maristella D"Addario, Manon

Amiguet, Isabel Fraccaroli, Andrea Ferretti, Julie Fluhr et Florie Delfour, qui ont contribué à

la collecte de données sur le comportement des balbuzards et l"abondance des poissons lors des transects marins en kayak, et ont rendu drôle le difficile travail de terrain en Corse. Merci

à ma FIAT Panda, courageuse aide de terrain depuis 2006, et à mon kayak “tranche de

citron" - Lemon King - qui, abandonné depuis 8 ans dans un jardin, est revenu surfer les vagues de la mer avec les thons et les dauphins. • Balearic Islands = Ostia que Bueno!! Rafel Triay gave me hospitality in the Balearics for ringing and equipping juvenile ospreys, in June 2013. The mission was perfectly organized thanks also to the support of the "Conselleria d'Agricultura, Medi Ambient i Territori (Govern de les Illes Balears)" in the persons of Juan Carlos Malmierca, Martí Mayol and Xavier Manzano. Antoni Muñoz Navarro kindly 10 provided important GPS data of adult ospreys. Félix de Pablo (the man of the kites) was of great help during fieldwork in Menorca. With Rafel I visited the most dangerous osprey nest of my life (until now): on a wonderful rocky cliff in Menorca Island with a superb view, the nest hosted the most aggressive female osprey “La Valiente" I"ve never met; she attacked us repeated times before allowing us to ring her chicks. Thanks to not have hit us so that I can still remember such moment and tell how brave can be a Spanish female osprey during the breeding season. Finally, I was lucky enough to experience the atmosphere of La Festa de Sant Joan de Ciutadella de Menorca during which inhabitants of the village opened their own doors and invited me to drink and sing in great company....“ostia que bueno". • Morocco = Inchallah !

Il est étonnant de noter que les coutumes des habitants de la Méditerranée peuvent être

similaires malgré les grandes différences de traditions, d"histoires et de langues entre les pays.

Je remercie les bons amis que j"ai rencontrés dans la région de Rif et du Parc National Al Hoceima, qui ont rejoint avec enthousiasme la " mission-balbuzard marocaine » : Houssine Nibani, Hamid Rguibi Idrissi, Mohamed Jabran, Abdennadi Abarkach, Abdelaziz El Edrissi, Abdeljebbar Qninba, Abdelouaed Kaikai, Sarrah El Hadri et Hamed El Fateh. Leur grande

habileté à résoudre les imprévus durant le travail de terrain nous ont permis de mener à bien la

mission. Les grimpeurs des nids difficiles à Al Hoceima étaient Jean Marie, Rémy Choquet et Rafel Triay. Pierre-Christian Beaubrun m"a fourni des données utiles et d"importants conseils pratiques. Je remercie aussi les pécheurs locaux qui nous ont emmenés le long des côtes du

parc, et l"âne nommé “chargé de mission", qui a porté le poids de la mission sur ses épaules

en 2014. Je remercie également Fabrice Bernard et Sébastien Renou, les coordinateurs du Conservatoire du Littoral qui ont coordonné le programme PIM-Albatros, pour avoir financé

les missions au Maroc. Pendant ces missions j"ai été accompagné de mon collègue du

Conservatoire du Littoral Mathieu Thévenet, qui a été indispensable à l"accomplissement de

ces missions et est devenu un très bon ami en France (un tarpin des fêtes). J"espère que nous

continuerons à partager de bons moments et des projets dans les années à venir. • Montpellier

Dans le cadre de la cotutelle j"ai été accueilli au Centre d"Ecologie Fonctionnelle et Evolutive

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