Using individual-based bioenergetic models to predict the aggregate effects of disturbance on populations: A case study with beaked whales and Navy sonar.

Anthropogenic activities can lead to changes in animal behavior. Predicting population consequences of these behavioral changes requires integrating short-term individual responses into models that forecast population dynamics across multiple generations. This is especially challenging for long-lived animals, because of the different time scales involved. Beaked whales are a group of deep-diving odontocete whales that respond behaviorally when exposed to military mid-frequency active sonar (MFAS), but the effect of these nonlethal responses on beaked whale populations is unknown. Population consequences of aggregate exposure to MFAS was assessed for two beaked whale populations that are regularly present on U.S. Navy training ranges where MFAS is frequently used. Our approach integrates a wide range of data sources, including telemetry data, information on spatial variation in habitat quality, passive acoustic data on the temporal pattern of sonar use and its relationship to beaked whale foraging activity, into an individual-based model with a dynamic bioenergetic module that governs individual life history. The predicted effect of disturbance from MFAS on population abundance ranged between population extinction to a slight increase in population abundance. These effects were driven by the interaction between the temporal pattern of MFAS use, baseline movement patterns, the spatial distribution of prey, the nature of beaked whale behavioral response to MFAS and the top-down impact of whale foraging on prey abundance. Based on these findings, we provide recommendations for monitoring of marine mammal populations and highlight key uncertainties to help guide future directions for assessing population impacts of nonlethal disturbance for these and other long-lived animals.

The effect of two 12 kHz multibeam mapping surveys on the foraging behavior of Cuvier's beaked whales off of southern California.

The impact of multibeam echosounder (MBES) operations on marine mammals has been less studied compared to military sonars. To contribute to the growing body of MBES knowledge, echolocation clicks of foraging Cuvier's beaked whales were detected on the Southern California Antisubmarine Warfare Range (SOAR) hydrophones during two MBES surveys and assembled into foraging events called group vocal periods (GVPs). Four GVP characteristics were analyzed Before, During, and After 12 kHz MBES surveys at the SOAR in 2017 and 2019 to assess differences in foraging behavior with respect to the mapping activity. The number of GVP per hour increased During and After MBES surveys compared with Before. There were no other differences between non-MBES and MBES periods for the three other characteristics: the number of clicks per GVP, GVP duration, and click rate. These results indicate that there was not a consistent change in foraging behavior during the MBES surveys that would suggest a clear response. The animals did not leave the range nor stop foraging during MBES activity. These results are in stark contrast to those of analogous studies assessing the effect of Naval mid-frequency active sonar on beaked whale foraging, where beaked whales stopped echolocating and left the area.

Beaked whale (Mesoplodon densirostris) passive acoustic detection in increasing ambient noise.

Passive acoustic detection is being increasingly used to monitor visually cryptic cetaceans such as Blainville's beaked whales (Mesoplodon densirostris) that may be especially sensitive to underwater sound. The efficacy of passive acoustic detection is traditionally characterized by the probability of detecting the animal's sound emissions as a function of signal-to-noise ratio. The probability of detection can be predicted using accepted, but not necessarily accurate, models of the underwater acoustic environment. Recent field studies combining far-field hydrophone arrays with on-animal acoustic recording tags have yielded the location and time of each sound emission from tagged animals, enabling in-situ measurements of the probability of detection. However, tagging studies can only take place in calm seas and so do not reflect the full range of ambient noise conditions under which passive acoustic detection may be used. Increased surface-generated noise from wind and wave interaction degrades the signal-to-noise ratio of animal sound receptions at a given distance leading to a reduction in probability of detection. This paper presents a case study simulating the effect of increasing ambient noise on detection of M. densirostris foraging clicks recorded from a tagged whale swimming in the vicinity of a deep-water, bottom-mounted hydrophone array.

Beaked whales respond to simulated and actual navy sonar.

Beaked whales have mass stranded during some naval sonar exercises, but the cause is unknown. They are difficult to sight but can reliably be detected by listening for echolocation clicks produced during deep foraging dives. Listening for these clicks, we documented Blainville's beaked whales, Mesoplodon densirostris, in a naval underwater range where sonars are in regular use near Andros Island, Bahamas. An array of bottom-mounted hydrophones can detect beaked whales when they click anywhere within the range. We used two complementary methods to investigate behavioral responses of beaked whales to sonar: an opportunistic approach that monitored whale responses to multi-day naval exercises involving tactical mid-frequency sonars, and an experimental approach using playbacks of simulated sonar and control sounds to whales tagged with a device that records sound, movement, and orientation. Here we show that in both exposure conditions beaked whales stopped echolocating during deep foraging dives and moved away. During actual sonar exercises, beaked whales were primarily detected near the periphery of the range, on average 16 km away from the sonar transmissions. Once the exercise stopped, beaked whales gradually filled in the center of the range over 2-3 days. A satellite tagged whale moved outside the range during an exercise, returning over 2-3 days post-exercise. The experimental approach used tags to measure acoustic exposure and behavioral reactions of beaked whales to one controlled exposure each of simulated military sonar, killer whale calls, and band-limited noise. The beaked whales reacted to these three sound playbacks at sound pressure levels below 142 dB re 1 µPa by stopping echolocation followed by unusually long and slow ascents from their foraging dives. The combined results indicate similar disruption of foraging behavior and avoidance by beaked whales in the two different contexts, at exposures well below those used by regulators to define disturbance.

Estimating cetacean population density using fixed passive acoustic sensors: an example with Blainville's beaked whales.

Methods are developed for estimating the size/density of cetacean populations using data from a set of fixed passive acoustic sensors. The methods convert the number of detected acoustic cues into animal density by accounting for (i) the probability of detecting cues, (ii) the rate at which animals produce cues, and (iii) the proportion of false positive detections. Additional information is often required for estimation of these quantities, for example, from an acoustic tag applied to a sample of animals. Methods are illustrated with a case study: estimation of Blainville's beaked whale density over a 6 day period in spring 2005, using an 82 hydrophone wide-baseline array located in the Tongue of the Ocean, Bahamas. To estimate the required quantities, additional data are used from digital acoustic tags, attached to five whales over 21 deep dives, where cues recorded on some of the dives are associated with those received on the fixed hydrophones. Estimated density was 25.3 or 22.5 animals/1000 km(2), depending on assumptions about false positive detections, with 95% confidence intervals 17.3-36.9 and 15.4-32.9. These methods are potentially applicable to a wide variety of marine and terrestrial species that are hard to survey using conventional visual methods.