Multi-colony approaches to study migratory and foraging strategies in pelagic seabirds

Movement is a widespread characteristic in the animal kingdom —occurring at many spatiotemporal scales— with consequences at an individual, population, species, and 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.El moviment és una característica omnipresent en el regne animal, a les més diverses escales espacio-temporals i amb conseqüències a diferents nivells (individual,. poblacional, específic 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. Els resultats d’aquesta tesi tenen rellevància per a la ecologia del moviment en general, ja que són aplicables a dades de moviment individual de qualsevol espècie, independentment de l’aparell de seguiment utilitzat, i en el camp de l’ecologia de les aus marines, ja que proporciona nous coneixements sobre els diferents factors afectant la distribució i l’ús de l’espai, tant durant la cria com durant la hivernada, en aus pelàgiques