The Umwelt concept of Jakob von Uexküll considers animals as subjects that inhabit their own subjective universe which is determined by the animal's sensory perception and cognitive abilities. Dolphins present an interesting species to investigate its Umwelt because cetaceans underwent a drastic change in lifestyle in the course of evolution because these mammals returned from a terrestrial life back into the water. Although bottlenose dolphins are intensively studied there are still many knowledge gaps. Here we studied some aspects of the dolphins' Umwelt by asking: (1) how their nocturnal acoustic Umwelt is arranged; (2) what the production of vocal copies can tell us about the dolphins' perception of their environment; (3) whether they are able to perceive tastes (4) or odours; (5) whether they are sensitive to magnetic stimuli. We found that the dolphins' nocturnal Umwelt was characterized by a temporally patterned vocal activity that also included vocal copies of sounds that the dolphins had heard during the day. This is a striking separation between auditory memory formation and vocal copy production and the copies might be a vocally expressed nocturnal rehearsal of day events. Thus, vocalizations can serve as possible indicators of events or objects that are meaningful to the dolphins. Regarding dolphins' perceptive abilities, we found that they were sensitive to both gustatory and olfactory food-related stimuli. They might use this ability to locate and/or evaluate prey. Further, dolphins responded to a magnetic stimulus, suggesting that they are magnetosensitive, what could be useful for navigation. So far, chemo- and magnetoreception have not been considered seriously as potentially functional in dolphins. The results obtained during this thesis fill some of the gaps that still exist in the knowledge of the dolphin's Umwelt and therefore contribute to a better understanding of this species. Moreover, they illustrate that even already intensively studied species may still hold important facets of their biology to reveal and that research should broaden the view and remain unbiased when studying a topic.

Le milieu marin est un espace difficile à étudier et notre connaissance des cétacés et des oiseaux marins présente encore de nombreuses lacunes, notamment concernant leur distribution au large. Le milieu marin pélagique est caractérisé par une forte variabilité spatiotemporelle, les différents processus écologiques composant cet écosystème étant très dynamiques dans le temps et l'espace. Il est donc crucial pour ces espèces marines d'anticiper ces variations afin de se maintenir dans le milieu. Grâce à des campagnes en mer à grandes échelles réalisées en Manche, Golfe de Gascogne et Méditerranée occidentale, cette thèse vise à améliorer la connaissance de la distribution des cétacés et oiseaux marins dans ces régions, et explorer l'impact de la variabilité spatiotemporelle de l'océan sur leur écologie. Ce travail se focalise notamment sur deux échelles temporelles, les variations saisonnières et interannuelles. Dans un premier temps, nous étudierons les variations de préférences d'habitats des espèces étudiées à ces deux échelles. Les cétacés et oiseaux marins étant des espèces longévives caractérisées par de longues périodes d'apprentissage au début de leur vie, nous explorerons également comment la réponse à la saisonnalité varie en fonction de l'expérience chez une espèce commune dans la région, le fou de Bassan. Enfin, nous finirons par explorer les liens prédateurs-proies dans le Golfe de Gascogne, en considérant notamment l'accessibilité de ces proies en termes de taille et de profondeur. Ainsi, cette thèse nous permettra de discuter les implications de ce travail concernant l'écologie des cétacés et oiseaux marins, ainsi que leur conservation. / Marine ecosystem is hard to study and several gaps remain in our understanding of cetaceans and seabirds, particularly regarding their at-sea distributions. The pelagic ecosystem is characterised by an important spatiotemporal variability, thanks to the space and time dynamics of its constituting ecological processes. Top predators thus have to anticipate this variability to sustain themselves in the ecosystem. Based on large-scale surveys across waters of the English Channel, Bay of Biscay and north-western Mediterranean Sea, this thesis aimed at improving knowledge regarding the distribution of cetaceans and seabirds in the area, and to explore the impact of ocean spatiotemporal variability on their ecology. This work focused on two temporal scales, the seasonal and interannual variability. We first explored the variations of habitat preferences of studied species at these two temporal scales. Cetaceans and seabirds being long-lived species characterised by long learning periods in their early life, the response to seasonality was investigated according to age and experience in northern gannets, the most abundant seabird species in the area. Finally, the predator-prey association within the Bay of Biscay was also explored, by contrasting prey accessibility in terms of size and depth. The implications of this work regarding the ecology of marine top predators, as well as their conservation, are finally discussed.

Effective management of the potential environmental impacts of naval sonar requires quantitative data on the behaviour and hearing physiology of cetaceans. Here, novel experimental and analytical methods were used to obtain such information and to test the effectiveness of an operational mitigation method for naval sonar. A Bayesian method was developed to estimate whale locations through time, integrating visual observations with measurements from on-animal inertial, acoustic, depth, and Fastloc-GPS sensors. The track reconstruction method was applied to 13 humpback whale (Megaptera novaeangliae) data sets collected during a multi-disciplinary behavioural response study in Norwegian waters. Thirty-one controlled exposure experiments with and without active transmissions of 1.3-2 kHz sounds were conducted using a moving vessel that towed a sonar source. Dose-response functions, representing the relationships between measured sonar dose and behavioural responses identified from the reconstructed tracks, predicted that 50% of the humpbacks would initiate avoidance at a relatively high received sound pressure level of 166 dB re 1 µPa. Very similar dose-response functions were obtained for cessation of feeding. In a laboratory study, behavioural reaction times of a harbour porpoise (Phocoena phocoena) to sonar-like sounds were measured using operant conditioning and a psychoacoustic method. Auditory weighting functions, which can be used to improve dose-response functions, were obtained for the porpoise based on the assumption that sounds of equal loudness elicit equal reaction time. Additional analyses of the humpback whale data set provided evidence that ramp-up of naval sonar mitigates harmful sound levels in responsive cetaceans located directly in the path of the source, and suggested that a subset of the humpback whale population, such as mother-calf pairs, and more responsive species would benefit from the use of sonar ramp-up. The findings in this thesis are intended to inform sound exposure criteria and mitigation guidelines for anthropogenic noise exposure to cetaceans.

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