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Funding was provided by the US Navy Living Marine Resources and Office of Naval Research programs, Netherlands Ministry of Defence, Norwegian Ministry of Defence, UK Defence Science and Technology Laboratory, and DGA French Ministry of Defence. Acoustic signals travel efficiently in the marine environment, allowing soniferous predators and prey to eavesdrop on each other. Our results with four cetacean species indicate that they use acoustic information to assess predation risk and have evolved mechanisms to reduce predation risk by ceasing foraging. Species that more readily gave up foraging in response to predatory sounds of killer whales also decreased foraging more during 1- to 4-kHz sonar exposures, indicating that species exhibiting costly antipredator responses also have stronger behavioral reactions to anthropogenic noise. This advance in our understanding of the drivers of disturbance helps us to predict what species and habitats are likely to be most severely impacted by underwater noise pollution in oceans undergoing increasing anthropogenic activities. As human activities impact virtually every animal habitat on the planet, identifying species at-risk from disturbance is a priority. Cetaceans are an example taxon where responsiveness to anthropogenic noise can be severe but highly species and context specific, with source–receiver characteristics such as hearing sensitivity only partially explaining this variability. Here, we predicted that ecoevolutionary factors that increase species responsiveness to predation risk also increase responsiveness to anthropogenic noise. We found that reductions in intense-foraging time during exposure to 1- to 4-kHz naval sonar and predatory killer whale sounds were highly correlated (r = 0.92) across four cetacean species. Northern bottlenose whales ceased foraging completely during killer whale and sonar exposures, followed by humpback, long-finned pilot, and sperm whales, which reduced intense foraging by 48 to 97level responses to killer whale playbacks, implying a similar level of perceived risk. The correlation cannot be solely explained by hearing sensitivity, indicating that species- and context-specific antipredator adaptations also shape cetacean responses to human-made noise. Species that are more responsive to predator presence are predicted to be more disturbance sensitive, implying a looming double whammy for Arctic cetaceans facing increased anthropogenic and predator activity with reduced ice cover. Publisher PDF

Funding: This work was funded by the US Office of Naval Research, grant number N00014-16-1-2858: ‘PCoD+: Developing widely-applicable models of the population consequences of disturbance’. Mike Dowd was supported by a Natural Sciences and Engineering Research Council of Canada Discovery grant. Understanding the impacts of anthropogenic sound on marine mammals is important for effective mitigation and management. Sound impacts can cause behavioral changes that lead to displacement from preferred habitat and can have negative influence on vital rates. Here, we develop a movement model to better understand and simulate how whales respond to anthropogenic sound over ecologically meaningful space and time scales. The stochastic model is based on a sequential Monte Carlo sampler (a particle filter). The movement model takes account of vertical dive information and is influenced by the underwater soundscape and the historical whale distribution in the region. In the absence of noise disturbance, the simulator is shown to recover the historical whale distribution in the region. When noise disturbance is incorporated, the whale’s behavioral response is determined through a dose–response function dependent on the received level of sound. The aggregate impact is assessed by considering both the duration of foraging loss and the spatial shift to alternate (and potentially less favorable) habitat. Persistence of the behavioral response in time is treated through a ‘disruption’ parameter. We apply the approach to a population of fin whales whose distribution overlaps naval sonar testing activities beside the Southern California range complex. The simulation shows the consequences of one year of naval sonar disturbance are a function of: i) how loud the sound source is, ii) where the disturbed whales are relative to preferred (high density) habitat, and iii) how long a whale takes before returning to a pre-disturbance state. The movement simulator developed here is a generic movement modeling tool that can be adapted for different species, different regions, and any acoustic disturbances with known impacts on animal populations. Publisher PDF

Research funding was provided by the US Office of Naval Research and the Ministries of Defence of Norway, the Netherlands and France as well as the UK Natural Environmental Research Council. A key issue when investigating effects of anthropogenic noise on cetacean behavior is to identify the biological significance of the responses. Predator presence can be considered a natural high-level disturbance stimulus to which prey animals have evolved adaptive response strategies to reduce their risk of predation by altering behavior away from fitness-enhancing activities such as foraging. By contrasting the type and magnitude (duration, severity, consistency) of behavioral responses to anthropogenic noise and playback of killer whale (KW) sounds that simulated predator presence, this study aimed to provide a relative index of the disturbance level as an indication of the biological significance of responses to the anthropogenic stimulus. Using multi-sensor tags as well as visual observations of surface behavior of adult male sperm whales, we assessed a comprehensive range of behavioral metrics that could reduce individuals’ fitness if altered for a biologically relevant duration. Combining previously published results and new analyses, we showed that the responses to 1-2 kHz upsweep naval sonar and to KW playback were very similar, including horizontal avoidance, interruption of foraging or resting activities and an increase in social sound production. However, only KW playbacks elicited grouping behaviors, indicating that this social response component was specific to predator detection. Animals responded to a lesser extent to 6-7 kHz upsweep naval sonar, indicating weaker disturbance effects. Our study demonstrates the benefit of using anti-predator responses as a reference of disturbance when evaluating the relative impacts of anthropogenic stimuli, which can be of particular interest in studies of threatened species such as sperm whales. Publisher PDF

The SOCAL project was funded by the US Navy Chief of Naval Operations Environmental Readiness Division and US Office of Naval Research. The 3S project was funded by the Norwegian Ministry of Defence, the US Office of Naval Research, the Netherlands Ministry of Defence and DGA French Ministry of Defence. The MOCHA project was funded by the US Office of Naval Research. Tyack received funding from the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland) and their support is gratefully acknowledged. MASTS is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions. Minke whales are difficult to study and little information exists regarding their responses to anthropogenic sound. This study pools data from behavioural response studies off California and Norway. Data are derived from four tagged animals, of which one from each location was exposed to naval sonar signals. Statistical analyses were conducted using Mahalanobis distance to compare overall changes in parameters summarising dive behaviour, avoidance behaviour, and potential energetic costs of disturbance. Our quantitative analysis showed that both animals initiated avoidance behaviour, but responses were not associated with unusual dive behaviour. In one exposed animal the avoidance of the sonar source included a 5-fold increase in horizontal speed away from the source, implying a significant increase in metabolic rate. Despite the different environmental settings and exposure contexts, clear changes in behaviour were observed providing the first insights into the nature of responses to human noise for this wide-ranging species. Postprint

Exposure to underwater sound can cause permanent hearing loss and other physiological effects in marine animals. To reduce this risk, naval sonars are sometimes gradually increased in intensity at the start of transmission (‘ramp-up’). Here, we conducted experiments in which tagged humpback whales were approached with a ship to test whether a sonar operation preceded by ramp-up reduced three risk indicators – maximum sound pressure level (SPLmax), cumulative sound exposure level (SELcum) and minimum source–whale range (Rmin) – compared with a sonar operation not preceded by ramp-up. Whales were subject to one no-sonar control session and either two successive ramp-up sessions (RampUp1, RampUp2) or a ramp-up session (RampUp1) and a full-power session (FullPower). Full-power sessions were conducted only twice; for other whales we used acoustic modelling that assumed transmission of the full-power sequence during their no-sonar control. Averaged over all whales, risk indicators in RampUp1 (n=11) differed significantly from those in FullPower (n=12) by −3.0 dB (SPLmax), −2.0 dB (SELcum) and +168 m (Rmin), but not significantly from those in RampUp2 (n=9). Only five whales in RampUp1, four whales in RampUp2 and none in FullPower or control sessions avoided the sound source. For RampUp1, we found statistically significant differences in risk indicators between whales that avoided the sonar and whales that did not: −4.7 dB (SPLmax), −3.4 dB (SELcum) and +291 m (Rmin). In contrast, for RampUp2, these differences were smaller and not significant. This study suggests that sonar ramp-up has a positive but limited mitigative effect for humpback whales overall, but that ramp-up can reduce the risk of harm more effectively in situations when animals are more responsive and likely to avoid the sonar, e.g. owing to novelty of the stimulus, when they are in the path of an approaching sonar ship., Highlighted Article: Variation in behavioural responses of humpback whales limits the effectiveness of a ramp-up procedure used to mitigate physiological effects of naval sonar on marine animals.

Minke whales are difficult to study and little information exists regarding their responses to anthropogenic sound. This study pools data from behavioural response studies off California and Norway. Data are derived from four tagged animals, of which one from each location was exposed to naval sonar signals. Statistical analyses were conducted using Mahalanobis distance to compare overall changes in parameters summarising dive behaviour, avoidance behaviour, and potential energetic costs of disturbance. Our quantitative analysis showed that both animals initiated avoidance behaviour, but responses were not associated with unusual dive behaviour. In one exposed animal the avoidance of the sonar source included a 5-fold increase in horizontal speed away from the source, implying a significant increase in metabolic rate. Despite the different environmental settings and exposure contexts, clear changes in behaviour were observed providing the first insights into the nature of responses to human noise for this wide-ranging species.

The authors would like to thank sponsors, NL Ministry of Defence, NOR Ministry of Defence, U.S. Office of Naval Research (N00014-08-1-0984, N00014-10-1-0355, N00014-14-1-0390), FR Ministry of Defence (DGA; public market no. 15860052), World Wildlife Fund Norway (9E0682), and French Total Foundation and Bleustein-Blanchet Foundation. The statistical development work was supported by a separate grant from the U.S. Office of Naval Research (N00014-12-1-0204), under the project entitled Multi-study OCean acoustics Human effects Analysis (MOCHA). Time allocation to different activities and habitats enables individuals to modulate their perceived risks and access to resources, and can reveal important trade-offs between fitness-enhancing activities (e.g., feeding vs. social behavior). Species with long reproductive cycles and high parental investment, such as marine mammals, rely on such behavioral plasticity to cope with rapid environmental change, including anthropogenic stressors. We quantified activity budgets of free-ranging long-finned pilot whales in order to assess individual time trade-offs between foraging and other behaviors in different individual and ecological contexts, and during experimental sound exposures. The experiments included 1-2 and 6-7 kHz naval sonar exposures (a potential anthropogenic stressor), playback of killer whale (a potential predator/competitor) vocalizations, and negative controls. We combined multiple time series data from digital acoustic recording tags (DTAG) as well as group-level social behavior data from visual observations of tagged whales at the surface. The data were classified into near-surface behaviors and dive types (using a hidden Markov model for dive transitions) and aggregated into time budgets. On average, individuals (N=19) spent most of their time (69%) resting and transiting near surface, 21% in shallow dives (depth

Exposure to underwater sound can cause permanent hearing loss and other physiological effects in marine animals. To reduce this risk, naval sonars are sometimes gradually increased in intensity at the start of transmission ('ramp-up'). Here, we conducted experiments in which tagged humpback whales were approached with a ship to test whether a sonar operation preceded by ramp-up reduced three risk indicators - maximum sound pressure level (SPLmax), cumulative sound exposure level (SELcum) and minimum source-whale range (Rmin) - compared with a sonar operation not preceded by ramp-up. Whales were subject to one no-sonar control session and either two successive ramp-up sessions (RampUp1, RampUp2) or a ramp-up session (RampUp1) and a full-power session (FullPower). Full-power sessions were conducted only twice; for other whales we used acoustic modelling that assumed transmission of the full-power sequence during their no-sonar control. Averaged overall whales, risk indicators in RampUp1 (n=11) differed significantly from those in FullPower (n=12) by -3.0 dB (SPLmax), -2.0 dB (SELcum) and +168 m (Rmin), but not significantly from those in RampUp2 (n=9). Only five whales in RampUp1, four whales in RampUp2 and none in FullPower or control sessions avoided the sound source. For RampUp1, we found statistically significant differences in risk indicators between whales that avoided the sonar and whales that did not: -4.7 dB (SPLmax), -3.4 dB (SELcum) and +291 m (Rmin). In contrast, for RampUp2, these differences were smaller and not significant. This study suggests that sonar ramp-up has a positive but limited mitigative effect for humpback whales overall, but that ramp-up can reduce the risk of harm more effectively in situations when animals are more responsive and likely to avoid the sonar, e.g. owing to novelty of the stimulus, when they are in the path of an approaching sonar ship. © 2017. Published by The Company of Biologists Ltd |.

Exposure to underwater sound can cause permanent hearing loss and other physiological effects in marine animals. To reduce this risk, naval sonars are sometimes gradually increased in intensity at the start of transmission (‘ramp-up’). Here, we conducted experiments in which tagged humpback whales were approached with a ship to test whether a sonar operation preceded by ramp-up reduced three risk indicators – maximum sound pressure level (SPLmax), cumulative sound exposure level (SELcum) and minimum source–whale range (Rmin) – compared with a sonar operation not preceded by ramp-up. Whales were subject to one no-sonar control session and either two successive ramp-up sessions (RampUp1, RampUp2) or a ramp-up session (RampUp1) and a full-power session (FullPower). Full-power sessions were conducted only twice; for other whales we used acoustic modelling that assumed transmission of the full-power sequence during their no-sonar control. Averaged over all whales, risk indicators in RampUp1 (n=11) differed significantly from those in FullPower (n=12) by −3.0 dB (SPLmax), −2.0 dB (SELcum) and +168 m (Rmin), but not significantly from those in RampUp2 (n=9). Only five whales in RampUp1, four whales in RampUp2 and none in FullPower or control sessions avoided the sound source. For RampUp1, we found statistically significant differences in risk indicators between whales that avoided the sonar and whales that did not: −4.7 dB (SPLmax), −3.4 dB (SELcum) and +291 m (Rmin). In contrast, for RampUp2, these differences were smaller and not significant. This study suggests that sonar ramp-up has a positive but limited mitigative effect for humpback whales overall, but that ramp-up can reduce the risk of harm more effectively in situations when animals are more responsive and likely to avoid the sonar, e.g. owing to novelty of the stimulus, when they are in the path of an approaching sonar ship., his work was supported by the Office of Naval Research [N00014-10-1-0355], The Netherlands Ministry of Defence, Norwegian Ministry of Defence and French Ministry of Defence. Additional support was provided through a PhD studentship withmatched funding from The Netherlands Ministry of Defence and the Natural Environment Research Council [NE/J500276/1 to P.J.W.]; and the MASTS (The Marine Alliance for Science and Technology for Scotland) pooling initiative to P.L.T. MASTS is funded by the Scottish Funding Council [HR09011] and contributing institutions.

Exposure to underwater sound can cause permanent hearing loss and other physiological effects in marine animals. To reduce this risk, naval sonars are sometimes gradually increased in intensity at the start of transmission ('ramp-up'). Here, we conducted experiments in which tagged humpback whales were approached with a ship to test whether a sonar operation preceded by ramp-up reduced three risk indicators - maximum sound pressure level (SPLmax), cumulative sound exposure level (SELcum) and minimum source-whale range (Rmin) - compared with a sonar operation not preceded by ramp-up. Whales were subject to one no-sonar control session and either two successive ramp-up sessions (RampUp1, RampUp2) or a ramp-up session (RampUp1) and a full-power session (FullPower). Full-power sessions were conducted only twice; for other whales we used acoustic modelling that assumed transmission of the full-power sequence during their no-sonar control. Averaged overall whales, risk indicators in RampUp1 (n=11) differed significantly from those in FullPower (n=12) by -3.0 dB (SPLmax), -2.0 dB (SELcum) and +168 m (Rmin), but not significantly from those in RampUp2 (n=9). Only five whales in RampUp1, four whales in RampUp2 and none in FullPower or control sessions avoided the sound source. For RampUp1, we found statistically significant differences in risk indicators between whales that avoided the sonar and whales that did not: -4.7 dB (SPLmax), -3.4 dB (SELcum) and +291 m (Rmin). In contrast, for RampUp2, these differences were smaller and not significant. This study suggests that sonar ramp-up has a positive but limited mitigative effect for humpback whales overall, but that ramp-up can reduce the risk of harm more effectively in situations when animals are more responsive and likely to avoid the sonar, e.g. owing to novelty of the stimulus, when they are in the path of an approaching sonar ship. © 2017. Published by The Company of Biologists Ltd |.

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. Postprint

We would like to thank 3S partners and funders especially for enabling this research (NL Ministry of Defence, NOR Ministry of Defence, US Office of Naval Research, and World Wildlife Fund, Norway). PLT was supported by the Scottish Funding Council (grant HR09011) through the Marine Alliance for Science and Technology for Scotland. The time and energetic costs of behavioral responses to incidental and experimental sonar exposures, as well as control stimuli, were quantified using hidden state analysis of time series of acoustic and movement data recorded by tags (DTAG) attached to 12 sperm whales (Physeter macrocephalus) using suction cups. Behavioral state transition modeling showed that tagged whales switched to a non-foraging, non-resting state during both experimental transmissions of low-frequency active sonar from an approaching vessel (LFAS; 1-2 kH z, source level 214 dB re 1 μPa m, four tag records) and playbacks of potential predator (killer whale, Orcinus orca) sounds broadcast at naturally occurring sound levels as a positive control from a drifting boat (five tag records). Time spent in foraging states and the probability of prey capture attempts were reduced during these two types of exposures with little change in overall locomotion activity, suggesting an effect on energy intake with no immediate compensation. Whales switched to the active non-foraging state over received sound pressure levels of 131-165 dB re 1 μPa during LFAS exposure. In contrast, no changes in foraging behavior were detected in response to experimental negative controls (no-sonar ship approach or noise control playback) or to experimental medium-frequency active sonar exposures (MFAS; 6-7 kH z, source level 199 re 1 μPa m, received sound pressure level [SPL] = 73-158 dB re 1 μPa). Similarly, there was no reduction in foraging effort for three whales exposed to incidental, unidentified 4.7-5.1 kH z sonar signals received at lower levels (SPL = 89-133 dB re 1 μPa). These results demonstrate that similar to predation risk, exposure to sonar can affect functional behaviors, and indicate that increased perception of risk with higher source level or lower frequency may modulate how sperm whales respond to anthropogenic sound. Publisher PDF

A key issue when investigating effects of anthropogenic noise on cetacean behavior is to identify the biological significance of the responses. Predator presence can be considered a natural high-level disturbance stimulus to which prey animals have evolved adaptive response strategies to reduce their risk of predation by altering behavior away from fitness-enhancing activities such as foraging. By contrasting the type and magnitude (duration, severity, consistency) of behavioral responses to anthropogenic noise and playback of killer whale (KW) sounds that simulated predator presence, this study aimed to provide a relative index of the disturbance level as an indication of the biological significance of responses to the anthropogenic stimulus. Using multi-sensor tags as well as visual observations of surface behavior of adult male sperm whales, we assessed a comprehensive range of behavioral metrics that could reduce individuals' fitness if altered for a biologically relevant duration. Combining previously published results and new analyses, we showed that the responses to 1-2 kHz upsweep naval sonar and to KW playback were very similar, including horizontal avoidance, interruption of foraging or resting activities and an increase in social sound production. However, only KW playbacks elicited grouping behaviors, indicating that this social response component was specific to predator detection. Animals responded to a lesser extent to 6-7 kHz upsweep naval sonar, indicating weaker disturbance effects. Our study demonstrates the benefit of using anti-predator responses as a reference of disturbance when evaluating the relative impacts of anthropogenic stimuli, which can be of particular interest in studies of threatened species such as sperm whales. © The authors 2016.

The time and energetic costs of behavioral responses to incidental and experimental sonar exposures, as well as control stimuli, were quantifi ed using hidden state analysis of time series of acoustic and movement data recorded by tags ( DTAG ) attached to 12 sperm whales ( Physeter macrocephalus ) using suction cups. Behavioral state transition modeling showed that tagged whales switched to a non- foraging, non- resting state during both experimental transmissions of low- frequency active sonar from an approaching vessel ( LFAS ; 1–2 kH z, source level 214 dB re 1 μPa m, four tag records) and playbacks of potential predator (killer whale, Orcinus orca ) sounds broadcast at naturally occurring sound levels as a positive control from a drifting boat (fi ve tag records). Time spent in foraging states and the probability of prey capture attempts were reduced during these two types of exposures with little change in overall locomotion activity, suggesting an effect on energy intake with no immediate compensation. Whales switched to the active non- foraging state over received sound pressure levels of 131–165 dB re 1 μPa during LFAS exposure. In contrast, no changes in foraging behavior were detected in response to experimental negative controls (no- sonar ship approach or noise control playback) or to experimental medium- frequency active sonar exposures ( MFAS ; 6–7 kH z, source level 199 re 1 μPa m, received sound pressure level [ SPL ] = 73–158 dB re 1 μPa). Similarly, there was no reduction in foraging effort for three whales exposed to incidental, unidentified 4.7–5.1 kH z sonar signals received at lower levels ( SPL = 89–133 dB re 1 μPa). These results demonstrate that similar to predation risk, exposure to sonar can affect functional behaviors, and indicate that increased perception of risk with higher source level or lower frequency may modulate how sperm whales respond to anthropogenic sound.

The time and energetic costs of behavioral responses to incidental and experimental sonar exposures, as well as control stimuli, were quantifi ed using hidden state analysis of time series of acoustic and movement data recorded by tags ( DTAG ) attached to 12 sperm whales ( Physeter macrocephalus ) using suction cups. Behavioral state transition modeling showed that tagged whales switched to a non- foraging, non- resting state during both experimental transmissions of low- frequency active sonar from an approaching vessel ( LFAS ; 1–2 kH z, source level 214 dB re 1 μPa m, four tag records) and playbacks of potential predator (killer whale, Orcinus orca ) sounds broadcast at naturally occurring sound levels as a positive control from a drifting boat (fi ve tag records). Time spent in foraging states and the probability of prey capture attempts were reduced during these two types of exposures with little change in overall locomotion activity, suggesting an effect on energy intake with no immediate compensation. Whales switched to the active non- foraging state over received sound pressure levels of 131–165 dB re 1 μPa during LFAS exposure. In contrast, no changes in foraging behavior were detected in response to experimental negative controls (no- sonar ship approach or noise control playback) or to experimental medium- frequency active sonar exposures ( MFAS ; 6–7 kH z, source level 199 re 1 μPa m, received sound pressure level [ SPL ] = 73–158 dB re 1 μPa). Similarly, there was no reduction in foraging effort for three whales exposed to incidental, unidentified 4.7–5.1 kH z sonar signals received at lower levels ( SPL = 89–133 dB re 1 μPa). These results demonstrate that similar to predation risk, exposure to sonar can affect functional behaviors, and indicate that increased perception of risk with higher source level or lower frequency may modulate how sperm whales respond to anthropogenic sound.

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. © 2016 The authors.

Contributions of P.M., P.T., C.C., S.D., F.V., P.W., L.M.L., T.N. and S.H. were funded by the US Office of Naval Research. Contributions of P.K., L.K. and L.S. were funded by the Norwegian Ministry of Defence. Contributions of F.L., S.v.IJ. and A.v.B. were funded by The Netherlands Ministry of Defence. Fieldwork contributions of L.M.L. and T.N. were funded by the Strategic Environmental Research and Development Program (SERDP). The analysis component of L.M.L.'s contribution was funded by DGA French Ministry of Defence. P.T. acknowledges the support of the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland) in the completion of this study. MASTS is funded by the Scottish Funding Council (grant no. HR09011) and contributing institutions. Although northern bottlenose whales were the most heavily hunted beaked whale, we have little information about this species in its remote habitat of the North Atlantic Ocean. Underwater anthropogenic noise and disruption of their natural habitat may be major threats, given the sensitivity of other beaked whales to such noise disturbance. We attached dataloggers to 13 northern bottlenose whales and compared their natural sounds and movements to those of one individual exposed to escalating levels of 1–2 kHz upsweep naval sonar signals. At a received sound pressure level (SPL) of 98 dB re 1 μPa, the whale turned to approach the sound source, but at a received SPL of 107 dB re 1 μPa, the whale began moving in an unusually straight course and then made a near 180° turn away from the source, and performed the longest and deepest dive (94 min, 2339 m) recorded for this species. Animal movement parameters differed significantly from baseline for more than 7 h until the tag fell off 33–36 km away. No clicks were emitted during the response period, indicating cessation of normal echolocation-based foraging. A sharp decline in both acoustic and visual detections of conspecifics after exposure suggests other whales in the area responded similarly. Though more data are needed, our results indicate high sensitivity of this species to acoustic disturbance, with consequent risk from marine industrialization and naval activity. Publisher PDF

Controlled exposure experiments using 1 to2 kHz sonar signals were conducted with 11 humpback whales (Megaptera novaeangliae), one minke whale (Balaenoptera acutorostrata), and one northern bottlenose whale (Hyperoodon ampullatus) during three field trials from 2011 to 2013. Ship approaches without sonar transmis-sions, playbacks of killer whale vocalizations, and broadband noise were conducted as controls. Behavioural parameters such as horizontal movement, diving, social interactions, and vocalizations were recorded by animal-attached tags and by visual and acoustic tracking. Based on these data, two expert panels independently scored the severity of behavioural changes that were judged likely to be responses to the experimental stimuli, using a severity scale ranging from no effect (0) to high potential to affect vital rates (9) if exposed repeatedly. After scoring, consensus was reached with a third-party moderator. In humpback whales, killer whale playbacks induced more severe responses than sonar exposure, and both sonar exposures and killer whale playbacks induced more responses and responses of higher severity than the no-sonar ship approaches and broadband noise playbacks. The most common response during sonar exposures in all three species was avoidance of the sound source. The most severe responses to sonar (severity 8) were progressive high-speed avoidance by the minke whale and long-term area avoidance by the bottlenose whale. Other severe responses included prolonged avoidance and cessation of feeding (severity 7). The minke whale and bottlenose whale started avoiding the source at a received sound pressure level (SPL) of 146 and 130 dB re 1 μPa, respectively. Humpback whales generally had less severe responses that were triggered at higher received levels. The probability of severity scores with the potential to affect vital rates increased with increasing sound exposure level (SEL). The single experiments with minke and bottlenose whales suggest they have greater susceptibility to sonar disturbance than humpback whales, but additional studies are needed to confirm this result.

Controlled exposure experiments using 1 to2 kHz sonar signals were conducted with 11 humpback whales (Megaptera novaeangliae), one minke whale (Balaenoptera acutorostrata), and one northern bottlenose whale (Hyperoodon ampullatus) during three field trials from 2011 to 2013. Ship approaches without sonar transmis-sions, playbacks of killer whale vocalizations, and broadband noise were conducted as controls. Behavioural parameters such as horizontal movement, diving, social interactions, and vocalizations were recorded by animal-attached tags and by visual and acoustic tracking. Based on these data, two expert panels independently scored the severity of behavioural changes that were judged likely to be responses to the experimental stimuli, using a severity scale ranging from no effect (0) to high potential to affect vital rates (9) if exposed repeatedly. After scoring, consensus was reached with a third-party moderator. In humpback whales, killer whale playbacks induced more severe responses than sonar exposure, and both sonar exposures and killer whale playbacks induced more responses and responses of higher severity than the no-sonar ship approaches and broadband noise playbacks. The most common response during sonar exposures in all three species was avoidance of the sound source. The most severe responses to sonar (severity 8) were progressive high-speed avoidance by the minke whale and long-term area avoidance by the bottlenose whale. Other severe responses included prolonged avoidance and cessation of feeding (severity 7). The minke whale and bottlenose whale started avoiding the source at a received sound pressure level (SPL) of 146 and 130 dB re 1 μPa, respectively. Humpback whales generally had less severe responses that were triggered at higher received levels. The probability of severity scores with the potential to affect vital rates increased with increasing sound exposure level (SEL). The single experiments with minke and bottlenose whales suggest they have greater susceptibility to sonar disturbance than humpback whales, but additional studies are needed to confirm this result.

Although northern bottlenose whales were the most heavily hunted beaked whale, we have little information about this species in its remote habitat of the North Atlantic Ocean. Underwater anthropogenic noise and disruption of their natural habitat may be major threats, given the sensitivity of other beaked whales to such noise disturbance. We attached dataloggers to 13 northern bottlenose whales and compared their natural sounds and movements to those of one individual exposed to escalating levels of 1–2 kHz upsweep naval sonar signals. At a received sound pressure level (SPL) of 98 dB re 1 μPa, the whale turned to approach the sound source, but at a received SPL of 107 dB re 1 μPa, the whale began moving in an unusually straight course and then made a near 180° turn away from the source, and performed the longest and deepest dive (94 min, 2339 m) recorded for this species. Animal movement parameters differed significantly from baseline for more than 7 h until the tag fell off 33–36 km away. No clicks were emitted during the response period, indicating cessation of normal echolocation-based foraging. A sharp decline in both acoustic and visual detections of conspecifics after exposure suggests other whales in the area responded similarly. Though more data are needed, our results indicate high sensitivity of this species to acoustic disturbance, with consequent risk from marine industrialization and naval activity.

There is increasing concern about the potential effects of noise pollution on marine life in the world’s oceans. For marine mammals, anthropogenic sounds may cause behavioral disruption, and this can be quantified using a risk function that relates sound exposure to a measured behavioral response. Beaked whales are a taxon of deep diving whales that may be particularly susceptible to naval sonar as the species has been associated with sonar-related mass stranding events. Here we derive the first empirical risk function for Blainville’s beaked whales (Mesoplodon densirostris) by combining in situ data from passive acoustic monitoring of animal vocalizations and navy sonar operations with precise ship tracks and sound field modeling. The hydrophone array at the Atlantic Undersea Test and Evaluation Center, Bahamas, was used to locate vocalizing groups of Blainville’s beaked whales and identify sonar transmissions before, during, and after Mid-Frequency Active (MFA) sonar operations. Sonar transmission times and source levels were combined with ship tracks using a sound propagation model to estimate the received level (RL) at each hydrophone. A generalized additive model was fitted to data to model the presence or absence of the start of foraging dives in 30-minute periods as a function of the corresponding sonar RL at the hydrophone closest to the center of each group. This model was then used to construct a risk function that can be used to estimate the probability of a behavioral change (cessation of foraging) the individual members of a Blainville’s beaked whale population might experience as a function of sonar RL. The function predicts a 0.5 probability of disturbance at a RL of 150dBrms re µPa (CI: 144 to 155) This is 15dB lower than the level used historically by the US Navy in their risk assessments but 10 dB higher than the current 140 dB step-function Publisher PDF

Naval anti-submarine sonars produce intense sounds within the hearing range of Atlantic herring (Clupea harengus). In this study, schools of Atlantic herring were exposed to sonar signals of 1–2 kHz (low-frequency active sonar, LFAS) and 6–7 kHz (mid-frequency active sonar, MFAS) and playbacks of killer whale feeding sounds during their summer feeding migration in the Norwegian Sea. The fish schools neither significantly dived nor changed their packing density in response to the LFAS and MFAS transmissions received by the fish at estimated sound pressure levels (SPLs; RMS) up to 176 and 157 dB re 1 mPa and estimated cumulative sound exposure levels up to 181 and 162 dB re 1 mPa2 s, respectively. In contrast, killer whale feeding sounds induced diving responses at received SPLs at -150 dB re 1 mPa. Herring behaviour was studied by using a 116-kHz hull mounted fishery sonar. This seems a promising method for studying the behaviour of free-ranging fish in situations in which other methods are difficult to use, such as migrating schools and fish close to the surface.

Naval anti-submarine sonars produce intense sounds within the hearing range of Atlantic herring (Clupea harengus). In this study, schools of Atlantic herring were exposed to sonar signals of 1–2 kHz (low-frequency active sonar, LFAS) and 6–7 kHz (mid-frequency active sonar, MFAS) and playbacks of killer whale feeding sounds during their summer feeding migration in the Norwegian Sea. The fish schools neither significantly dived nor changed their packing density in response to the LFAS and MFAS transmissions received by the fish at estimated sound pressure levels (SPLs; RMS) up to 176 and 157 dB re 1 mPa and estimated cumulative sound exposure levels up to 181 and 162 dB re 1 mPa2 s, respectively. In contrast, killer whale feeding sounds induced diving responses at received SPLs at -150 dB re 1 mPa. Herring behaviour was studied by using a 116-kHz hull mounted fishery sonar. This seems a promising method for studying the behaviour of free-ranging fish in situations in which other methods are difficult to use, such as migrating schools and fish close to the surface.