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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.

Behavioural responses can reveal important fitness trade-offs and ecological traps in evolutionarily novel contexts created by anthropogenic stimuli, and are of increasing conservation concern due to possible links to population-level impacts. This thesis illustrates the use of proxies for energy acquisition and expenditure within multivariate and state-based modelling approaches to quantify the relative time and energetic costs of behavioural disturbance for a deep-diving marine mammal (Physeter macrocephalus) in foraging grounds in Kaikoura Canyon (New Zealand) and near Lofoten Islands (Norway). A conceptual framework is first developed to identify and explore links between individual motivation, condition and external constraints to behavioural disturbance [Chapter 1]. The following chapters then use data from behavioural response studies (BRS) to: 1) derive biologically relevant metrics of behaviour [all chapters], 2) investigate effects of boat-based focal follows and tagging procedures [Chapters 2-3], and 3) relate responses to specific disturbance stimuli (distance, approach, noise) from whale-watching [Chapter 2], naval sonar and playback of presumed natural predator (killer whale Orcinus orca) sounds [Chapter 4]. A novel hidden state model was developed to estimate behavioural budgets of tagged sperm whales from multiple streams of biologging (DTAG) data [Chapter 3]. Sperm whales traded off time spent at foraging depths in a non-foraging and non-resting state in response to both tag boat presence, 1-2 kHz naval sonar (SPL 131-165 rms re 1μPa) and mammal-eating killer whale sound playbacks, indicating that parallel non-lethal costs were incurred in both anthropogenic disturbance and presumed antipredatory contexts. While behavioural responses were highly variable by individual, biologically informed state-based models appeared effective to control for variability in energy proxies across different functional contexts. These results and Chapter 5 “linking buzzes to prey” demonstrate that behavioural context is a signal that can aid understanding of how individual non-lethal disturbance responses can impact fitness.