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Acoustic sequences have been described in a range of species and in varying complexity. Cetaceans are known to produce complex song displays but these are generally limited to mysticetes; little is known about call combinations in odontocetes. Here we investigate call combinations produced by killer whales ( Orcinus orca ), a highly social and vocal species. Using acoustic recordings from 22 multisensor tags, we use a first order Markov model to show that transitions between call types or subtypes were significantly different from random, with repetitions and specific call combinations occurring more often than expected by chance. The mixed call combinations were composed of two or three calls and were part of three call combination clusters. Call combinations were recorded over several years, from different individuals, and several social clusters. The most common call combination cluster consisted of six call (sub-)types. Although different combinations were generated, there were clear rules regarding which were the first and last call types produced, and combinations were highly stereotyped. Two of the three call combination clusters were produced outside of feeding contexts, but their function remains unclear and further research is required to determine possible functions and whether these combinations could be behaviour- or group-specific.

Impact assessments for sonar operations typically use received sound levels to predict behavioural disturbance in marine mammals. However, there are indications that cetaceans may learn to associate exposures from distant sound sources with lower perceived risk. To investigate the roles of source distance and received level in an area without frequent sonar activity, we conducted multi-scale controlled exposure experiments ( n = 3) with 12 northern bottlenose whales near Jan Mayen, Norway. Animals were tagged with high-resolution archival tags ( n = 1 per experiment) or medium-resolution satellite tags ( n = 9 in total) and subsequently exposed to sonar. We also deployed bottom-moored recorders to acoustically monitor for whales in the exposed area. Tagged whales initiated avoidance of the sound source over a wide range of distances (0.8–28 km), with responses characteristic of beaked whales. Both onset and intensity of response were better predicted by received sound pressure level (SPL) than by source distance. Avoidance threshold SPLs estimated for each whale ranged from 117–126 dB re 1 µPa, comparable to those of other tagged beaked whales. In this pristine underwater acoustic environment, we found no indication that the source distances tested in our experiments modulated the behavioural effects of sonar, as has been suggested for locations where whales are frequently exposed to sonar.

Interactions are common among marine species which use sound as the primary form of communication, yet the role of acoustic signals in mediating these interactions remains poorly understood. Long-finned pilot whales ( Globicephala melas ) are attracted to killer whale ( Orcinus orca ) sounds, leading to antagonistic interactions. To test whether these interactions are acoustically mediated in both directions, playback experiments ( n  = 15) using pilot whale and control sound stimuli were conducted on eight killer whales equipped with multi-sensor tags. To assess behavioural responses to the playbacks, we applied hidden Markov models (HMMs) to movement and acoustic data, and fitted univariate regression models to a horizontal movement reaction score, calling rate, and group behaviour variables. The tagged whales exhibited an avoidance response to pilot whale sounds, evidenced by fast, directed movement away from the sound source and increased cohesion and alignment of group members. Calling rate often increased initially, followed by a pronounced decrease. These findings demonstrate that killer whales, the oceans’ apex predators, respond to acoustic signals of pilot whales and likely perceive their presence as a threat, similar to naval sonar. This study provides important insights into the complexity of cetacean behaviour and acoustic mechanisms shaping multi-species community dynamics.

Studying the baseline behavior of deep‐diving mammals can substantially improve our understanding of these species' ecology and provide important benchmarks to evaluate effects of changes in climate and anthropogenic activities. Despite being the most abundant beaked whale in the Arctic and subarctic, information on the behavior of northern bottlenose whales (Hyperoodon ampullatus) is limited. This study used records from 13 satellite tags deployed off Jan Mayen in June–July 2014–2016 to provide an extensive description of the dive behavior of Hyperoodon for the Nordic Seas. A total of 8372 dives, collected over 224 days (or 5376 h), were analyzed. The whales performed extreme dives of up to 2288 m deep and 98 min long—deeper and longer than previously reported for behavior in presumed undisturbed contexts. Individuals spent on average 18% of the time at depths shallower than 40 m, and 22%, 47%, and 12% in epi‐, meso‐, and bathypelagic dives, respectively. Epipelagic dives averaged 123 m (s.d.: 46 m) in depth and 11 min (5 min) in duration. Mesopelagic dives averaged 441 m (217 m) and 24 min (11 min) and were performed at a mean rate of 1.46 h−1. Bathypelagic dives averaged 1487 m (366 m) and 55 min (13 min) and were performed at a mean rate of 0.23 h−1. The distribution of dive depths was less bimodal than typically reported for other beaked whales, and all dive profiles contained periods of continuous, consecutive deep dives. Benthic diving occurred at meso‐ and especially bathypelagic depths and was individual specific, varying from 8% to 51% of the animal's bathypelagic dives. Overall, our findings demonstrate that northern bottlenose whales have extraordinary capabilities to dive, and presumably feed, throughout the water column including at the sea floor. High rates of deep dives highlight the importance of the Iceland and Norwegian Seas to this population of deep‐sea predators. This study used records from 13 satellite tags deployed off Jan Mayen to provide an extensive description of the dive behavior of northern bottlenose whales in the Nordic Seas. Our findings demonstrate that this species has extraordinary capabilities to dive, and presumably feed, throughout the water column including at the sea floor. High rates of deep dives highlight the importance of the Iceland and Norwegian Seas habitat to this population of deep‐sea predators.

Killer whales (KW) may be predators or competitors of other cetaceans. Since their foraging behavior and acoustics differ among populations (‘ecotypes’), we hypothesized that other cetaceans can eavesdrop on KW sounds and adjust their behavior according to the KW ecotype. We performed playback experiments on long-finned pilot whales (Globicephala melas) in Norway using familiar fish-eating KW sounds (fKW) simulating a sympatric population that might compete for foraging areas, unfamiliar mammal-eating KW sounds (mKW) simulating a potential predator threat, and two control sounds. We assessed behavioral responses using animal-borne multi-sensor tags and surface visual observations. Pilot whales barely changed behavior to a broadband noise (CTRL−), whereas they were attracted and exhibited spyhops to fKW, mKW, and to a repeated-tonal upsweep signal (CTRL+). Whales never stopped nor started feeding in response to fKW, whereas they reduced or stopped foraging to mKW and CTRL+. Moreover, pilot whales joined other subgroups in response to fKW and CTRL+, whereas they tightened individual spacing within group and reduced time at surface in response to mKW. Typical active intimidation behavior displayed to fKW might be an antipredator strategy to a known low-risk ecotype or alternatively a way of securing the habitat exploited by a heterospecific sympatric population. Cessation of feeding and more cohesive approach to mKW playbacks might reflect an antipredator behavior towards an unknown KW ecotype of potentially higher risk. We conclude that pilot whales are able to acoustically discriminate between familiar and unfamiliar KW ecotypes, enabling them to adjust their behavior according to the perceived disturbance type.

Background High-resolution sound and movement recording tags offer unprecedented insights into the fine-scale foraging behaviour of cetaceans, especially echolocating odontocetes, enabling the estimation of a series of foraging metrics. However, these tags are expensive, making them inaccessible to most researchers. Time-Depth Recorders (TDRs), which have been widely used to study diving and foraging behaviour of marine mammals, offer a more affordable alternative. Unfortunately, data collected by TDRs are bi-dimensional (time and depth only), so quantifying foraging effort from those data is challenging. Methods A predictive model of the foraging effort of sperm whales ( Physeter macrocephalus ) was developed to identify prey capture attempts (PCAs) from time-depth data. Data from high-resolution acoustic and movement recording tags deployed on 12 sperm whales were downsampled to 1 Hz to match the typical TDR sampling resolution and used to predict the number of buzzes (i.e., rapid series of echolocation clicks indicative of PCAs). Generalized linear mixed models were built for dive segments of different durations (30, 60, 180 and 300 s) using multiple dive metrics as potential predictors of PCAs. Results Average depth, variance of depth and variance of vertical velocity were the best predictors of the number of buzzes. Sensitivity analysis showed that models with segments of 180 s had the best overall predictive performance, with a good area under the curve value (0.78 ± 0.05), high sensitivity (0.93 ± 0.06) and high specificity (0.64 ± 0.14). Models using 180 s segments had a small difference between observed and predicted number of buzzes per dive, with a median of 4 buzzes, representing a difference in predicted buzzes of 30%. Conclusions These results demonstrate that it is possible to obtain a fine-scale, accurate index of sperm whale PCAs from time-depth data alone. This work helps leveraging the potential of time-depth data for studying the foraging ecology of sperm whales and the possibility of applying this approach to a wide range of echolocating cetaceans. The development of accurate foraging indices from low-cost, easily accessible TDR data would contribute to democratize this type of research, promote long-term studies of various species in several locations, and enable analyses of historical datasets to investigate changes in cetacean foraging activity.

Background Detailed information about animal location and movement is often crucial in studies of natural behaviour and how animals respond to anthropogenic activities. Dead-reckoning can be used to infer such detailed information, but without additional positional data this method results in uncertainty that grows with time. Combining dead-reckoning with new Fastloc-GPS technology should provide good opportunities for reconstructing georeferenced fine-scale tracks, and should be particularly useful for marine animals that spend most of their time under water. We developed a computationally efficient, Bayesian state-space modelling technique to estimate humpback whale locations through time, integrating dead-reckoning using on-animal sensors with measurements of whale locations using on-animal Fastloc-GPS and visual observations. Positional observation models were based upon error measurements made during calibrations. Results High-resolution 3-dimensional movement tracks were produced for 13 whales using a simple process model in which errors caused by water current movements, non-location sensor errors, and other dead-reckoning errors were accumulated into a combined error term. Positional uncertainty quantified by the track reconstruction model was much greater for tracks with visual positions and few or no GPS positions, indicating a strong benefit to using Fastloc-GPS for track reconstruction. Compared to tracks derived only from position fixes, the inclusion of dead-reckoning data greatly improved the level of detail in the reconstructed tracks of humpback whales. Using cross-validation, a clear improvement in the predictability of out-of-set Fastloc-GPS data was observed compared to more conventional track reconstruction methods. Fastloc-GPS observation errors during calibrations were found to vary by number of GPS satellites received and by orthogonal dimension analysed; visual observation errors varied most by distance to the whale.  Conclusions By systematically accounting for the observation errors in the position fixes, our model provides a quantitative estimate of location uncertainty that can be appropriately incorporated into analyses of animal movement. This generic method has potential application for a wide range of marine animal species and data recording systems.

Monitoring the body condition of free-ranging marine mammals at different life-history stages is essential to understand their ecology as they must accumulate sufficient energy reserves for survival and reproduction. However, assessing body condition in free-ranging marine mammals is challenging. We cross-validated two independent approaches to estimate the body condition of humpback whales (Megaptera novaeangliae) at two feeding grounds in Canada and Norway: animal-borne tags (n = 59) and aerial photogrammetry (n = 55). Whales that had a large length-standardized projected area in overhead images (i.e. whales looked fatter) had lower estimated tissue body density (TBD) (greater lipid stores) from tag data. Linking both measurements in a Bayesian hierarchical model to estimate the true underlying (hidden) tissue body density (uTBD), we found uTBD was lower (−3.5 kg m−3) in pregnant females compared to adult males and resting females, while in lactating females it was higher (+6.0 kg m−3). Whales were more negatively buoyant (+5.0 kg m−3) in Norway than Canada during the early feeding season, possibly owing to a longer migration from breeding areas. While uTBD decreased over the feeding season across life-history traits, whale tissues remained negatively buoyant (1035.3 ± 3.8 kg m−3) in the late feeding season. This study adds confidence to the effectiveness of these independent methods to estimate the body condition of free-ranging whales.

Controlled exposure experiments (CEEs) have demonstrated that naval pulsed active sonar (PAS) can induce costly behavioral responses in cetaceans similar to antipredator responses. New generation continuous active sonars (CAS) emit lower amplitude levels but more continuous signals. We conducted CEEs with PAS, CAS and no-sonar control on free-ranging sperm whales in Norway. Two panels blind to experimental conditions concurrently inspected acoustic-and-movement-tag data and visual observations of tagged whales and used an established severity scale (0–9) to assign scores to putative responses. Only half of the exposures elicited a response, indicating overall low responsiveness in sperm whales. Responding whales (10 of 12) showed more, and more severe responses to sonar compared to no-sonar. Moreover, the probability of response increased when whales were previously exposed to presence of predatory and/or competing killer or long-finned pilot whales. Various behavioral change types occurred over a broad range of severities (1–6) during CAS and PAS. When combining all behavioral types, the proportion of responses to CAS was significantly higher than no-sonar but not different from PAS. Responses potentially impacting vital rates i.e., with severity ≥4, were initiated at received cumulative sound exposure levels (dB re 1 μPa2 s) of 137–177 during CAS and 143–181 during PAS.

Modern long-range naval sonars are a potential disturbance for marine mammals and can cause disruption of feeding in cetaceans. We examined the lunge-feeding behaviour of humpback whales Megaptera novaeangliae before, during and after controlled exposure experiments with naval sonar by use of acoustic and motion sensor archival tags attached to each animal. Lunge-feeding by humpback whales entails a strong acceleration to increase speed before engulfing a large volume of prey-laden water, which can be identified by an acoustic signature characterized by a few seconds of high-level flow-noise followed by a rapid reduction, coinciding with a peak in animal acceleration. Over 2 successive seasons, 13 humpback whales were tagged. All were subject to a no-sonar control exposure, and 12 whales were exposed to 2 consecutive sonar exposure sessions, with 1 h between sessions. The first sonar session resulted in an average 68% reduction in lunge rate during exposure compared to pre-exposure, and this reduction was significantly greater than any changes observed during the no-sonar control. During the second sonar session, reduction in lunge rate was 66% during sonar exposure compared to the pre-exposure level, but was not significant compared to the no-sonar control, likely due to a larger inter-individual variability because some individuals appeared to have habituated whereas others had not. Our results indicate that naval sonars operating near humpback whale feeding grounds may lead to reduced foraging and negative impacts on energy balance.