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Multi-Colony Approaches to Study Migratory and Foraging Strategies in Pelagic Seabirds

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Multi-Colony Approaches to Study Migratory and Foraging Strategies in Pelagic Seabirds Multi-colony approaches to study migratory and foraging strategies in pelagic seabirds Virginia Morera Pujol Aquesta tesi doctoral està subjecta a la llicència Reconeixement- NoComercial 4.0. Espanya de Creative Commons. Esta tesis doctoral está sujeta a la licencia Reconocimiento - NoComercial 4.0. España de Creative Commons. This doctoral thesis is licensed under the Creative Commons Attribution-NonCommercial 4.0. Spain License. Facultat de Biologia Departament de Biologia Animal, Ecologia i Ciències Ambientals Programa de Doctorat en Biodiversitat (HDK04) Multi-colony approaches to study migratory and foraging strategies in pelagic seabirds Una aproximació multi-colònia a l’estudi de les estratègies migratòries i de recerca d’aliment en aus pelàgiques Memòria presentada per Virginia Morera Pujol per optar al grau de Doctora per la Universitat de Barcelona, Barcelona, Setembre de 2019 La doctoranda, El director de la tesi, El director i tutor de la tesi, Acknowledgments lgú (no recordo qui) em va dir en algun moment d’aquest any boig que fes els agraïments amb temps, que després a última Ahora tindria les neurones fregides i no sabria a qui he d’agrair què. Tant de bo li hagués fet cas, però fer les coses amb temps no serà mai el meu estil, així que sense ordre ni prioritat, aquí van els meus agraïments: El Raül i el Jacob han corregut aquest final de marató amb mi i això, en si, ja és una fita. Més enllà d’aquest tram final, no sé ni com comptar, molt menys agrair, les estones de “Raül, baixem a buscar un cafè” que s’han convertit en hores i hores de discussió i resolució de problemes, i sense les quals aquesta tesi no estaria ni la meitat de bé (i ni parlar de la maleïda consistència!). Ni la de vegades que un “Jacob, jo això no ho sé fer” ha rebut un “però segur que te’n surts” per resposta, i ves per on, la majoria tenies raó! Aquesta confiança cega, de vegades tan desesperant, m’ha empès fins a on som ara, gràcies! Teresa y Jose me enseñaron todo lo que necesitaba saber para arrancar en esta carrera de locos, y me siguen enseñando cada día coses nuevas. Sin ellos, no hubiese partido con la ventaja que me ha permitido llegar hasta aquí. Con ellos y con Zuzana, Fernanda, Marta, Laura y Leia hemos remado en el mismo barco, haciendo el camino mucho más agradable. Siempre hay algo que aprender de tus compañeros de viaje, y yo he aprendido de cada una de vosotras. Eli, Moisès, Jazel, y demás compañeros no-jacobinos de despacho y departamento, gracias por las risas, las horas de comer, las cervezas de viernes tarde y tantas otras cosas. Amigues i amics de fora del món de la ciència, però que m’heu acompanyat en aquest camí, fent-lo tan més fàcil que no sé com me n’hagués sortit sense vosaltres. Sou enterament responsables de que jo no hagi acabat parant boja durant aquests quatre anys i mig. Sou el contrapunt que necessito per relativitzar i veure les coses amb una perspectiva difícil de guanyar quan et capfiques tant en un projecte, i mai ho podré agrair prou. En aquest últim tram de bogeria, de fet, no hagués menjat, ni vestit, ni probablement sobreviscut sense l’Alba i l’Emma cuidant-me, cuinant per a mi, recollint-me la roba estesa, i en general, tenint una paciència enorme amb la meva invasió de la casa per treballar a totes hores. The three months I spent in Saint Andrews have been one of the highlights of this already amazing journey. And they wouldn’t have been half as good without everyone at CREEM being so incredibly nice to me, without Claudia and Valentin being so welcoming and friendly, and without Andy and his immense patience and kindness in explaining to me what I’m sure were the most basic statistical concepts. Of course, I am also incredibly grateful to Janine for accepting me there, for teaching me so, so much, and for always encouraging me to learn more. Lastly, a big, huge thank you to Lucía, for being as awesome as she always was and for introducing me to her group of friends from the very beginning of my time there. Por último, a mi familia. No hay como agradecer una vida de apoyo a esta carrera loca que he escogido, de paciencia y comprensión con las ausencias por trabajo de campo, estancias, congresos y demás. Ha sido un largo camino hasta aquí, y lo que queda por delante, pero vaya donde vaya, siempre me dais un motivo para volver. Abstract ovement is a widespread characteristic in the animal kingdom —occurring at many spatiotemporal scales— with consequences at an individual, population, species, Mand even ecosystem level. It is a very diverse character, with many different drivers that stem from the way in which individuals interact with their environment. Of these, one of the most important is the distribution of resources, particularly for migratory and foraging movements. In migration, the search for an optimal environment involves movement at large spatiotemporal scales, following seasonal changes in resource distribution. In foraging movements, the search for resources happens at small spatiotemporal scales, and involves different strategies to optimise the search and capture of food, including the ability to obtain foraging cues from conspecifics. In seabirds, movement —at large and local scales— has deep repercussions in their life-history traits, evolutionary history, morphology, physiology and behaviour, which makes them a very valuable study group to understand the role, the causes and consequences of migratory and foraging movements in the ecology of marine top-predators. The study of migratory and foraging movements has been revolutionised by the development of smaller, cheaper and better tracking devices, promoting multi-colony, population and even species approaches to the study of animal movement, but which also come with a set of methodological challenges that have to be addressed in order to make unbiased inferences of space and habitat use at population or species level from individual movement data. In this thesis, we develop methods to test the possible biases introduced by the use of individual tracking data to infer distribution at a population or species level. we then apply these tools to a multi-colony dataset of non-breeding locations of Cory’s (Calonectris borealis), Scopoli’s (C. diomedea) and Cape Verde (C. edwardsii) shearwaters, to study their migratory connectivity and non-breeding habitat segregation at the colony, population and species level. Lastly, we apply state-of- the-art spatial models to study foraging distributions of three neighbouring colonies of Cory’s shearwaters, detect the segregation among them and unravel the environmental and behavioural drivers of this segregation. I developed several functions in the R environment aimed at the detection of the effects of individual site fidelity and temporal variability in the inference of spatial use at a colony or population level, and to calculate the degree in which the movements of a single population can be representative of those of the entire species. These tools are applicable to individual movement data regardless of the species or tracking device. we also used these tools to demonstrate the spatial and ecological segregation between the non-breeding distributions of three taxa of Calonectris shearwaters studied, as well as detecting a stronger degree of migratory connectivity at a population than at a colony level, indicating that individuals of different colonies within a population mix in the non-breeding areas, but birds from different populations do not, which has important implications for their population dynamics and for their conservation and management. Lastly, we demonstrated segregation among the foraging distributions of three neighbouring colonies of Cory’s shearwaters, both in the waters surrounding the colony and in distant, foraging grounds, finding evidences of both environmental and behavioural drivers behind this segregation, and suggesting a mechanism through which transfer of information between individuals can be shaping the distributions of foraging seabirds. The results of this thesis provide relevant tools for the field of movement ecology, as they can be used for analysing movements of mobile species, regardless of species, tracking device or spatiotemporal scale. In addition, they are relevant for the field of seabird ecology as they provide insights into the causes of space and habitat use in long-ranging pelagic seabirds. Resum l moviment és una característica omnipresent en el regne animal, a les més diverses escales espacio-temporals i amb conseqüències a diferents nivells (individual,. poblacional, Eespecífic i fins i tot ecosistèmic). La força impulsora del moviment més predominant i generalitzada és distribució dels recursos en el medi, important tant a gran escala (moviments migratoris) com en els moviments diaris de recerca d’aliment a escala petita. En les aus marines en particular, el moviment té repercussions profundes en les seves característiques morfològiques i de comportament, en la seva història de vida, i la seva història evolutiva. El desenvolupament de dispositius de seguiment més petits, barats i precisos ha promocionat la proliferació d’estudis del moviment animal des d’un punt de vista multi-colònia, de població i fins i tot d’espècie. En aquesta tesi, desenvolupo diferents funcions per testar els biaixos introduïts en l’estudi del moviment, a través de dades de seguiment individual, a nivell de població o espècie. Posteriorment, utilitzo aquestes eines per a analitzar la connectivitat migratòria i la segregació dels hàbitats d’hivernada, des d’un punt de vista multi-colònia, de les baldrigues cendroses de l’Atlàntic (Calonectris borealis), del Mediterrani (C. diomedea), i de Cap Verd (C. edwardsii). Per últim, aplico un mètode innovador de modelatge espacial per a estudiar les distribucions d’alimentació de baldrigues cendroses de l’Atlàntic criant en tres colònies veïnes, per detectar-ne la segregació i descobrir-ne les causes, tant ambientals com comportamentals, incloent com els diferents mecanismes de transferència d’informació entre individus poden afectar a aquestes distribucions.
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