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.

Comparison of fin whale 20 Hz call detections by deep-water mobile autonomous and stationary recorders.

Acoustically equipped deep-water mobile autonomous platforms can be used to survey for marine mammals over intermediate spatiotemporal scales. Direct comparisons to fixed recorders are necessary to evaluate these tools as passive acoustic monitoring platforms. One glider and two drifting deep-water floats were simultaneously deployed within a deep-water cabled hydrophone array to quantitatively assess their survey capabilities. The glider was able to follow a pre-defined track while float movement was somewhat unpredictable. Fin whale (Balaenoptera physalus) 20 Hz pulses were recorded by all hydrophones throughout the two-week deployment. Calls were identified using a template detector, which performed similarly across recorder types. The glider data contained up to 78% fewer detections per hour due to increased low-frequency flow noise present during glider descents. The glider performed comparably to the floats and fixed recorders at coarser temporal scales; hourly and daily presence of detections did not vary by recorder type. Flow noise was related to glider speed through water and dive state. Glider speeds through water of 25 cm/s or less are suggested to minimize flow noise and the importance of glider ballasting, detector characterization, and normalization by effort when interpreting glider-collected data and applying it to marine mammal density estimation are discussed.

Behavioral responses of individual blue whales (Balaenoptera musculus) to mid-frequency military sonar.

This study measured the degree of behavioral responses in blue whales (Balaenoptera musculus) to controlled noise exposure off the southern California coast. High-resolution movement and passive acoustic data were obtained from non-invasive archival tags (n=42) whereas surface positions were obtained with visual focal follows. Controlled exposure experiments (CEEs) were used to obtain direct behavioral measurements before, during and after simulated and operational military mid-frequency active sonar (MFAS), pseudorandom noise (PRN) and controls (no noise exposure). For a subset of deep-feeding animals (n=21), active acoustic measurements of prey were obtained and used as contextual covariates in response analyses. To investigate potential behavioral changes within individuals as a function of controlled noise exposure conditions, two parallel analyses of time-series data for selected behavioral parameters (e.g. diving, horizontal movement and feeding) were conducted. This included expert scoring of responses according to a specified behavioral severity rating paradigm and quantitative change-point analyses using Mahalanobis distance statistics. Both methods identified clear changes in some conditions. More than 50% of blue whales in deep-feeding states responded during CEEs, whereas no changes in behavior were identified in shallow-feeding blue whales. Overall, responses were generally brief, of low to moderate severity, and highly dependent on exposure context such as behavioral state, source-to-whale horizontal range and prey availability. Response probability did not follow a simple exposure-response model based on received exposure level. These results, in combination with additional analytical methods to investigate different aspects of potential responses within and among individuals, provide a comprehensive evaluation of how free-ranging blue whales responded to mid-frequency military sonar.

Understanding the population consequences of disturbance.

Managing the nonlethal effects of disturbance on wildlife populations has been a long-term goal for decision makers, managers, and ecologists, and assessment of these effects is currently required by European Union and United States legislation. However, robust assessment of these effects is challenging. The management of human activities that have nonlethal effects on wildlife is a specific example of a fundamental ecological problem: how to understand the population-level consequences of changes in the behavior or physiology of individual animals that are caused by external stressors. In this study, we review recent applications of a conceptual framework for assessing and predicting these consequences for marine mammal populations. We explore the range of models that can be used to formalize the approach and we identify critical research gaps. We also provide a decision tree that can be used to select the most appropriate model structure given the available data. Synthesis and applications: The implementation of this framework has moved the focus of discussion of the management of nonlethal disturbances on marine mammal populations away from a rhetorical debate about defining negligible impact and toward a quantitative understanding of long-term population-level effects. Here we demonstrate the framework's general applicability to other marine and terrestrial systems and show how it can support integrated modeling of the proximate and ultimate mechanisms that regulate trait-mediated, indirect interactions in ecological communities, that is, the nonconsumptive effects of a predator or stressor on a species' behavior, physiology, or life history.

Diving behaviour of Cuvier's beaked whales exposed to two types of military sonar.

Cuvier's beaked whales (Ziphius cavirostris) have stranded in association with mid-frequency active sonar (MFAS) use, and though the causative mechanism linking these events remains unclear, it is believed to be behaviourally mediated. To determine whether MFAS use was associated with behavioural changes in this species, satellite tags were used to record the diving and movements of 16 Cuvier's beaked whales for up to 88 days in a region of frequent MFAS training off the coast of Southern California. Tag data were combined with summarized records of concurrent bouts of high-power, surface-ship and mid-power, helicopter-deployed MFAS use, along with other potential covariates, in generalized additive mixed-effects models. Deep dives, shallow dives and surface intervals tended to become longer during MFAS use, with some variation associated with the total amount of overlapping MFAS during the behaviour. These changes in dives and surface intervals contributed to a longer interval between deep dives, a proxy for foraging disruption in this species. Most responses intensified with proximity and were more pronounced during mid-power than high-power MFAS use at comparable distances within approximately 50 km, despite the significantly lower source level of mid-power MFAS. However, distance-mediated responses to high-power MFAS, and increased deep dive intervals during mid-power MFAS, were evident up to approximately 100 km away.

Acoustic detection of biosonar activity of deep diving odontocetes at Josephine Seamount High Seas Marine Protected Area.

The temporal occurrence of deep diving cetaceans in the Josephine Seamount High Seas Marine Protected Area (JSHSMPA), south-west Portugal, was monitored using a passive acoustic recorder. The recorder was deployed on 13 May 2010 at a depth of 814 m during the North Atlantic Treaty Organization Centre for Maritime Research and Experimentation cruise "Sirena10" and recovered on 6 June 2010. The recorder was programmed to record 40 s of data every 2 min. Acoustic data analysis, for the detection and classification of echolocation clicks, was performed using automatic detector/classification systems: M3R (Marine Mammal Monitoring on Navy Ranges), a custom matlab program, and an operator-supervised custom matlab program to assess the classification performance of the detector/classification systems. M3R CS-SVM algorithm contains templates to detect beaked whales, sperm whales, blackfish (pilot and false killer whales), and Risso's dolphins. The detections of each group of odontocetes was monitored as a function of time. Blackfish and Risso's dolphins were detected every day, while beaked whales and sperm whales were detected almost every day. The hourly distribution of detections reveals that blackfish and Risso's dolphins were more active at night, while beaked whales and sperm whales were more active during daylight hours.

First long-term behavioral records from Cuvier's beaked whales (Ziphius cavirostris) reveal record-breaking dives.

Cuvier's beaked whales (Ziphius cavirostris) are known as extreme divers, though behavioral data from this difficult-to-study species have been limited. They are also the species most often stranded in association with Mid-Frequency Active (MFA) sonar use, a relationship that remains poorly understood. We used satellite-linked tags to record the diving behavior and locations of eight Ziphius off the Southern California coast for periods up to three months. The effort resulted in 3732 hr of dive data with associated regional movements--the first dataset of its kind for any beaked whale--and included dives to 2992 m depth and lasting 137.5 min, both new mammalian dive records. Deep dives had a group mean depth of 1401 m (s.d. = 137.8, n = 1142) and duration of 67.4 min (s.d. = 6.9). The group mean time between deep dives was 102.3 min (s.d. = 30.8, n = 783). While the previously described stereotypic pattern of deep and shallow dives was apparent, there was considerable inter- and intra-individual variability in most parameters. There was significant diel behavioral variation, including increased time near the surface and decreased shallow diving at night. However, maximum depth and the proportion of time spent on deep dives (presumed foraging), varied little from day to night. Surprisingly, tagged whales were present within an MFA sonar training range for 38% of days locations were received, and though comprehensive records of sonar use during tag deployments were not available, we discuss the effects frequent acoustic disturbance may have had on the observed behaviors. These data better characterize the true behavioral range of this species, and suggest caution should be exercised when drawing conclusions about behavior using short-term datasets.

A risk function for behavioral disruption of Blainville's beaked whales (Mesoplodon densirostris) from mid-frequency active sonar.

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 150 dBrms 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.

Presence and seasonal variation of deep diving foraging odontocetes around Kauai, Hawaii using remote autonomous acoustic recorders.

Ecological acoustic recorders (EARs) were moored off the bottom in relatively deep depths (609-710 m) at five locations around the island of Kauai. Initially, the EARs had an analog-to-digital sample rate of 64 kHz with 30-s recordings every 5 min. After the second deployment the sampling rate was increased to 80 kHz in order to better record beaked whale biosonar signals. The results of the 80 kHz recording are discussed in this manuscript and are the results of three deployments over a year's period (January 2010 to January 2011). Five categories of the biosonar signal detection of deep diving odontocetes were created, short-finned pilot whales, sperm whales, beaked whales, Risso's dolphins, and unknown dolphins. During any given day, at least one species of these deep diving odontocetes were detected. On many days, several species were detected. The biosonar signals of short-finned pilot whales were detected the most often with approximately 30% of all the signals, followed by beaked and sperm whales approximately 22% and 21% of all clicks, respectively. The seasonal patterns were not very strong except in the SW location with distinct peak in detection during the months of April-June 2010 period.

First direct measurements of behavioural responses by Cuvier's beaked whales to mid-frequency active sonar.

Most marine mammal- strandings coincident with naval sonar exercises have involved Cuvier's beaked whales (Ziphius cavirostris). We recorded animal movement and acoustic data on two tagged Ziphius and obtained the first direct measurements of behavioural responses of this species to mid-frequency active (MFA) sonar signals. Each recording included a 30-min playback (one 1.6-s simulated MFA sonar signal repeated every 25 s); one whale was also incidentally exposed to MFA sonar from distant naval exercises. Whales responded strongly to playbacks at low received levels (RLs; 89-127 dB re 1 µPa): after ceasing normal fluking and echolocation, they swam rapidly, silently away, extending both dive duration and subsequent non-foraging interval. Distant sonar exercises (78-106 dB re 1 µPa) did not elicit such responses, suggesting that context may moderate reactions. The observed responses to playback occurred at RLs well below current regulatory thresholds; equivalent responses to operational sonars could elevate stranding risk and reduce foraging efficiency.

Blue whales respond to simulated mid-frequency military sonar.

Mid-frequency military (1-10 kHz) sonars have been associated with lethal mass strandings of deep-diving toothed whales, but the effects on endangered baleen whale species are virtually unknown. Here, we used controlled exposure experiments with simulated military sonar and other mid-frequency sounds to measure behavioural responses of tagged blue whales (Balaenoptera musculus) in feeding areas within the Southern California Bight. Despite using source levels orders of magnitude below some operational military systems, our results demonstrate that mid-frequency sound can significantly affect blue whale behaviour, especially during deep feeding modes. When a response occurred, behavioural changes varied widely from cessation of deep feeding to increased swimming speed and directed travel away from the sound source. The variability of these behavioural responses was largely influenced by a complex interaction of behavioural state, the type of mid-frequency sound and received sound level. Sonar-induced disruption of feeding and displacement from high-quality prey patches could have significant and previously undocumented impacts on baleen whale foraging ecology, individual fitness and population health.

Using energetic models to investigate the survival and reproduction of beaked whales (family Ziphiidae).

Mass stranding of several species of beaked whales (family Ziphiidae) associated with exposure to anthropogenic sounds has raised concern for the conservation of these species. However, little is known about the species' life histories, prey or habitat requirements. Without this knowledge, it becomes difficult to assess the effects of anthropogenic sound, since there is no way to determine whether the disturbance is impacting the species' physical or environmental requirements. Here we take a bioenergetics approach to address this gap in our knowledge, as the elusive, deep-diving nature of beaked whales has made it hard to study these effects directly. We develop a model for Ziphiidae linking feeding energetics to the species' requirements for survival and reproduction, since these life history traits would be the most likely to be impacted by non-lethal disturbances. Our models suggest that beaked whale reproduction requires energy dense prey, and that poor resource availability would lead to an extension of the inter-calving interval. Further, given current information, it seems that some beaked whale species require relatively high quality habitat in order to meet their requirements for survival and reproduction. As a result, even a small non-lethal disturbance that results in displacement of whales from preferred habitats could potentially impact a population if a significant proportion of that population was affected. We explored the impact of varying ecological parameters and model assumptions on survival and reproduction, and find that calf and fetus survival appear more readily affected than the survival of adult females.

Nighttime foraging by deep diving echolocating odontocetes off the Hawaiian islands of Kauai and Ni'ihau as determined by passive acoustic monitors.

Remote autonomous ecological acoustic recorders (EARs) were deployed in deep waters at five locations around the island of Kauai and one in waters off Ni'ihau in the main Hawaiian island chain. The EARs were moored to the bottom at depths between 400 and 800 m. The data acquisition sampling rate was 80 kHz and acoustic signals were recorded for 30 s every 5 min to conserve battery power and disk space. The acoustic data were analyzed with the M3R (Marine Mammal Monitoring on Navy Ranges) software, an energy-ratio-mapping algorithm developed at Oregon State University and custom MATLAB programs. A variety of deep diving odontocetes, including pilot whales, Risso's dolphins, sperm whales, spinner and pan-tropical spotted dolphins, and beaked whales were detected at all sites. Foraging activity typically began to increase after dusk, peaked in the middle of the night and began to decrease toward dawn. Between 70% and 84% of biosonar clicks were detected at night. At present it is not clear why some of the known deep diving species, such as sperm whales and beaked whales, concentrate their foraging efforts at night.

Effective beam pattern of the Blainville's beaked whale (Mesoplodon densirostris) and implications for passive acoustic monitoring.

The presence of beaked whales in mass-strandings coincident with navy maneuvers has prompted the development of methods to detect these cryptic animals. Blainville's beaked whales, Mesoplodon densirostris, produce distinctive echolocation clicks during long foraging dives making passive acoustic detection a possibility. However, performance of passive acoustic monitoring depends upon the source level, beam pattern, and clicking behavior of the whales. In this study, clicks recorded from Digital acoustic Tags (DTags) attached to four M. densirostris were linked to simultaneous recordings from an 82-hydrophone bottom-mounted array to derive the source level and beam pattern of the clicks, as steps towards estimating their detectability. The mean estimated on-axis apparent source level for the four whales was 201 dBrms97. The mean 3 dB beamwidth and directivity index, estimated from sequences of clicks directed towards the far-field hydrophones, were 13° and 23 dB, respectively. While searching for prey, Blainville's beaked whales scan their heads horizontally at a mean rate of 3.6°/s over an angular range of some +/-10°. Thus, while the DI indicates a narrow beam, the area of ensonification over a complete foraging dive is large given the combined effects of body and head movements associated with foraging.

Estimating animal population density using passive acoustics.

Reliable estimation of the size or density of wild animal populations is very important for effective wildlife management, conservation and ecology. Currently, the most widely used methods for obtaining such estimates involve either sighting animals from transect lines or some form of capture-recapture on marked or uniquely identifiable individuals. However, many species are difficult to sight, and cannot be easily marked or recaptured. Some of these species produce readily identifiable sounds, providing an opportunity to use passive acoustic data to estimate animal density. In addition, even for species for which other visually based methods are feasible, passive acoustic methods offer the potential for greater detection ranges in some environments (e.g. underwater or in dense forest), and hence potentially better precision. Automated data collection means that surveys can take place at times and in places where it would be too expensive or dangerous to send human observers. Here, we present an overview of animal density estimation using passive acoustic data, a relatively new and fast-developing field. We review the types of data and methodological approaches currently available to researchers and we provide a framework for acoustics-based density estimation, illustrated with examples from real-world case studies. We mention moving sensor platforms (e.g. towed acoustics), but then focus on methods involving sensors at fixed locations, particularly hydrophones to survey marine mammals, as acoustic-based density estimation research to date has been concentrated in this area. Primary among these are methods based on distance sampling and spatially explicit capture-recapture. The methods are also applicable to other aquatic and terrestrial sound-producing taxa. We conclude that, despite being in its infancy, density estimation based on passive acoustic data likely will become an important method for surveying a number of diverse taxa, such as sea mammals, fish, birds, amphibians, and insects, especially in situations where inferences are required over long periods of time. There is considerable work ahead, with several potentially fruitful research areas, including the development of (i) hardware and software for data acquisition, (ii) efficient, calibrated, automated detection and classification systems, and (iii) statistical approaches optimized for this application. Further, survey design will need to be developed, and research is needed on the acoustic behaviour of target species. Fundamental research on vocalization rates and group sizes, and the relation between these and other factors such as season or behaviour state, is critical. Evaluation of the methods under known density scenarios will be important for empirically validating the approaches presented here.

Vessel noise affects beaked whale behavior: results of a dedicated acoustic response study.

Some beaked whale species are susceptible to the detrimental effects of anthropogenic noise. Most studies have concentrated on the effects of military sonar, but other forms of acoustic disturbance (e.g. shipping noise) may disrupt behavior. An experiment involving the exposure of target whale groups to intense vessel-generated noise tested how these exposures influenced the foraging behavior of Blainville's beaked whales (Mesoplodon densirostris) in the Tongue of the Ocean (Bahamas). A military array of bottom-mounted hydrophones was used to measure the response based upon changes in the spatial and temporal pattern of vocalizations. The archived acoustic data were used to compute metrics of the echolocation-based foraging behavior for 16 targeted groups, 10 groups further away on the range, and 26 non-exposed groups. The duration of foraging bouts was not significantly affected by the exposure. Changes in the hydrophone over which the group was most frequently detected occurred as the animals moved around within a foraging bout, and their number was significantly less the closer the whales were to the sound source. Non-exposed groups also had significantly more changes in the primary hydrophone than exposed groups irrespective of distance. Our results suggested that broadband ship noise caused a significant change in beaked whale behavior up to at least 5.2 kilometers away from the vessel. The observed change could potentially correspond to a restriction in the movement of groups, a period of more directional travel, a reduction in the number of individuals clicking within the group, or a response to changes in prey movement.

Cetacean population density estimation from single fixed sensors using passive acoustics.

Passive acoustic methods are increasingly being used to estimate animal population density. Most density estimation methods are based on estimates of the probability of detecting calls as functions of distance. Typically these are obtained using receivers capable of localizing calls or from studies of tagged animals. However, both approaches are expensive to implement. The approach described here uses a MonteCarlo model to estimate the probability of detecting calls from single sensors. The passive sonar equation is used to predict signal-to-noise ratios (SNRs) of received clicks, which are then combined with a detector characterization that predicts probability of detection as a function of SNR. Input distributions for source level, beam pattern, and whale depth are obtained from the literature. Acoustic propagation modeling is used to estimate transmission loss. Other inputs for density estimation are call rate, obtained from the literature, and false positive rate, obtained from manual analysis of a data sample. The method is applied to estimate density of Blainville's beaked whales over a 6-day period around a single hydrophone located in the Tongue of the Ocean, Bahamas. Results are consistent with those from previous analyses, which use additional tag data.

The relationship among oceanography, prey fields, and beaked whale foraging habitat in the Tongue of the Ocean.

Beaked whales, specifically Blainville's (Mesoplodon densirostris) and Cuvier's (Ziphius cavirostris), are known to feed in the Tongue of the Ocean, Bahamas. These whales can be reliably detected and often localized within the Atlantic Undersea Test and Evaluation Center (AUTEC) acoustic sensor system. The AUTEC range is a regularly spaced bottom mounted hydrophone array covering >350 nm(2) providing a valuable network to record anthropogenic noise and marine mammal vocalizations. Assessments of the potential risks of noise exposure to beaked whales have historically occurred in the absence of information about the physical and biological environments in which these animals are distributed. In the fall of 2008, we used a downward looking 38 kHz SIMRAD EK60 echosounder to measure prey scattering layers concurrent with fine scale turbulence measurements from an autonomous turbulence profiler. Using an 8 km, 4-leaf clover sampling pattern, we completed a total of 7.5 repeat surveys with concurrently measured physical and biological oceanographic parameters, so as to examine the spatiotemporal scales and relationships among turbulence levels, biological scattering layers, and beaked whale foraging activity. We found a strong correlation among increased prey density and ocean vertical structure relative to increased click densities. Understanding the habitats of these whales and their utilization patterns will improve future models of beaked whale habitat as well as allowing more comprehensive assessments of exposure risk to anthropogenic sound.