Modeling potential masking of echolocating sperm whales exposed to continuous 1-2 kHz naval sonar.

Modern active sonar systems can (almost) continuously transmit and receive sound, which can lead to more masking of important sounds for marine mammals than conventional pulsed sonar systems transmitting at a much lower duty cycle. This study investigated the potential of 1-2 kHz active sonar to mask echolocation-based foraging of sperm whales by modeling their echolocation detection process. Continuous masking for an echolocating sperm whale facing a sonar was predicted for sonar sound pressure levels of 160 dB re 1 µPa2, with intermittent masking at levels of 120 dB re 1 µPa2, but model predictions strongly depended on the animal orientation, harmonic content of the sonar, click source level, and target strength of the prey. The masking model predicted lower masking potential of buzz clicks compared to regular clicks, even though the energy source level is much lower. For buzz clicks, the lower source level is compensated for by the reduced two-way propagation loss to nearby prey during buzzes. These results help to predict what types of behavioral changes could indicate masking in the wild. Several key knowledge gaps related to masking potential of sonar in echolocating odontocetes were identified that require further investigation to assess the significance of masking.

Predicting acoustic dose associated with marine mammal behavioural responses to sound as detected with fixed acoustic recorders and satellite tags.

To understand the consequences of underwater noise exposure for cetaceans, there is a need for assessments of behavioural responses over increased spatial and temporal scales. Bottom-moored acoustic recorders and satellite tags provide such long-term and large spatial coverage of behaviour compared to short-duration acoustic-recording tags. However, these tools result in a decreased resolution of data from which an animal response can be inferred, and no direct recording of the sound received at the animal. This study discusses the consequence of the decreased resolution of data from satellite tags and fixed acoustic recorders on the acoustic dose estimated by propagation modelling and presents a method for estimating the range of sound levels that animals observed with these methods have received. This problem is illustrated using experimental results obtained during controlled exposures of northern bottlenose whales (Hyperoodon ampullatus) exposed to naval sonar, carried out near Jan Mayen, Norway. It is shown that variability and uncertainties in the sound field, resulting from limited sampling of the acoustic environment, as well as decreased resolution in animal locations, can lead to quantifiable uncertainties in the estimated acoustic dose associated with the behavioural response (in this case avoidance and cessation of foraging).

First indications that northern bottlenose whales are sensitive to behavioural disturbance from anthropogenic noise.

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.

High thresholds for avoidance of sonar by free-ranging long-finned pilot whales (Globicephala melas).

The potential effects of exposing marine mammals to military sonar is a current concern. Dose-response relationships are useful for predicting potential environmental impacts of specific operations. To reveal behavioral response thresholds of exposure to sonar, we conducted 18 exposure/control approaches to 6 long-finned pilot whales. Source level and proximity of sonar transmitting one of two frequency bands (1-2 kHz and 6-7 kHz) were increased during exposure sessions. The 2-dimensional movement tracks were analyzed using a changepoint method to identify the avoidance response thresholds which were used to estimate dose-response relationships. No support for an effect of sonar frequency or previous exposures on the probability of response was found. Estimated response thresholds at which 50% of population show avoidance (SPLmax=170 dB re 1 µPa, SELcum=173 dB re 1 µPa(2) s) were higher than previously found for other cetaceans. The US Navy currently uses a generic dose-response relationship to predict the responses of cetaceans to naval active sonar, which has been found to underestimate behavioural impacts on killer whales and beaked whales. The navy curve appears to match more closely our results with long-finned pilot whales, though it might underestimate the probability of avoidance for pilot-whales at long distances from sonar sources.

Changes in dive behavior during naval sonar exposure in killer whales, long-finned pilot whales, and sperm whales.

Anthropogenic underwater sound in the environment might potentially affect the behavior of marine mammals enough to have an impact on their reproduction and survival. Diving behavior of four killer whales (Orcinus orca), seven long-finned pilot whales (Globicephala melas), and four sperm whales (Physeter macrocephalus) was studied during controlled exposures to naval sonar [low frequency active sonar (LFAS): 1-2 kHz and mid frequency active sonar (MFAS): 6-7 kHz] during three field seasons (2006-2009). Diving behavior was monitored before, during and after sonar exposure using an archival tag placed on the animal with suction cups. The tag recorded the animal's vertical movement, and additional data on horizontal movement and vocalizations were used to determine behavioral modes. Killer whales that were conducting deep dives at sonar onset changed abruptly to shallow diving (ShD) during LFAS, while killer whales conducting deep dives at the onset of MFAS did not alter dive mode. When in ShD mode at sonar onset, killer whales did not change their diving behavior. Pilot and sperm whales performed normal deep dives (NDD) during MFAS exposure. During LFAS exposures, long-finned pilot whales mostly performed fewer deep dives and some sperm whales performed shallower and shorter dives. Acoustic recording data presented previously indicates that deep diving (DD) is associated with feeding. Therefore, the observed changes in dive behavior of the three species could potentially reduce the foraging efficiency of the affected animals.

Estimated Tissue and Blood N(2) Levels and Risk of Decompression Sickness in Deep-, Intermediate-, and Shallow-Diving Toothed Whales during Exposure to Naval Sonar.

Naval sonar has been accused of causing whale stranding by a mechanism which increases formation of tissue N(2) gas bubbles. Increased tissue and blood N(2) levels, and thereby increased risk of decompression sickness (DCS), is thought to result from changes in behavior or physiological responses during diving. Previous theoretical studies have used hypothetical sonar-induced changes in both behavior and physiology to model blood and tissue N(2) tension [Formula: see text], but this is the first attempt to estimate the changes during actual behavioral responses to sonar. We used an existing mathematical model to estimate blood and tissue N(2) tension [Formula: see text] from dive data recorded from sperm, killer, long-finned pilot, Blainville's beaked, and Cuvier's beaked whales before and during exposure to Low- (1-2 kHz) and Mid- (2-7 kHz) frequency active sonar. Our objectives were: (1) to determine if differences in dive behavior affects risk of bubble formation, and if (2) behavioral- or (3) physiological responses to sonar are plausible risk factors. Our results suggest that all species have natural high N(2) levels, with deep diving generally resulting in higher end-dive [Formula: see text] as compared with shallow diving. Sonar exposure caused some changes in dive behavior in both killer whales, pilot whales and beaked whales, but this did not lead to any increased risk of DCS. However, in three of eight exposure session with sperm whales, the animal changed to shallower diving, and in all these cases this seem to result in an increased risk of DCS, although risk was still within the normal risk range of this species. When a hypothetical removal of the normal dive response (bradycardia and peripheral vasoconstriction), was added to the behavioral response during model simulations, this led to an increased variance in the estimated end-dive N(2) levels, but no consistent change of risk. In conclusion, we cannot rule out the possibility that a combination of behavioral and physiological responses to sonar have the potential to alter the blood and tissue end-dive N(2) tension to levels which could cause DCS and formation of in vivo bubbles, but the actually observed behavioral responses of cetaceans to sonar in our study, do not imply any significantly increased risk of DCS.

Deadly diving? Physiological and behavioural management of decompression stress in diving mammals.

Decompression sickness (DCS; 'the bends') is a disease associated with gas uptake at pressure. The basic pathology and cause are relatively well known to human divers. Breath-hold diving marine mammals were thought to be relatively immune to DCS owing to multiple anatomical, physiological and behavioural adaptations that reduce nitrogen gas (N(2)) loading during dives. However, recent observations have shown that gas bubbles may form and tissue injury may occur in marine mammals under certain circumstances. Gas kinetic models based on measured time-depth profiles further suggest the potential occurrence of high blood and tissue N(2) tensions. We review evidence for gas-bubble incidence in marine mammal tissues and discuss the theory behind gas loading and bubble formation. We suggest that diving mammals vary their physiological responses according to multiple stressors, and that the perspective on marine mammal diving physiology should change from simply minimizing N(2) loading to management of the N(2) load. This suggests several avenues for further study, ranging from the effects of gas bubbles at molecular, cellular and organ function levels, to comparative studies relating the presence/absence of gas bubbles to diving behaviour. Technological advances in imaging and remote instrumentation are likely to advance this field in coming years.

Cutaneous heat flux models do not reliably predict metabolic rates of marine mammals.

Heat flux models have been used to predict metabolic rates of marine mammals, generally by estimating conductive heat transfer through their blubber layer. Recently, Kvadsheim et al. (1997) found that such models tend to overestimate metabolic rates, and that such errors probably result from the asymmetrical distribution of blubber. This problem may be avoided if reliable estimates of heat flux through the skin of the animals are obtained by using models that combine calculations of conductive heat flux through the skin and fur, and convective heat flux from the surface of the animal to the environment. We evaluated this approach based on simultaneous measurements of metabolic rates and of input parameters necessary for heat flux calculations, as obtained from four harp seals (Phoca groenlandica) resting in cold water. Heat flux estimates were made using two free convection models (double-flat-plate and cylindrical geometry) and one forced convection model (single-flat-plate geometry). We found that heat flux estimates generally underestimated metabolic rates, on average by 26-58%, and that small variations in input parameters caused large variations in these estimates. We conclude that cutaneous heat flux models are too inaccurate and sensitive to small errors in input parameters to provide reliable estimates of metabolic rates of marine mammals.

Blubber and flipper heat transfer in harp seals.

The trunk of marine mammals is encased in a blubber layer which provides thermal insulation that can be changed by circulatory adjustments. The extremities, on the other hand, are poorly insulated but have vascular arrangements constructed for prevention or promotion of heat loss depending on the thermal state of the animal. We have studied the importance of different body parts as sites for heat dissipation and also assessed the effect of circulatory adjustments on heat transfer through blubber, by combining direct measurements of heat flux from the flippers and trunk with simultaneous recordings of temperature gradients through the blubber and metabolic rates of harp seals (Phoca groenlandica) subjected to water temperatures between 1 and 24 degrees C. We also determined the thermal conductivity of blubber samples from the same animals after death, and compared this with the insulative properties of live blubber. At the lowest water temperatures, the insulative properties of live blubber were similar to those of dead blubber, and heat loss from the flippers only accounted for 2-6% of the metabolic heat production. As heat load increased with increasing water temperatures, the fraction of heat lost from the flippers increased, to 19-48% at 24 degrees C, while the fraction lost from the trunk decreased, despite an increase in the convective (circulatory) heat transfer through the blubber layer.