Communication in Cook Inlet beluga whales: Describing the vocal repertoire and masking of calls by commercial ship noise.

Many species rely on acoustic communication to coordinate activities and communicate to conspecifics. Cataloging vocal behavior is a first step towards understanding how individuals communicate information and how communication may be degraded by anthropogenic noise. The Cook Inlet beluga population is endangered with an estimated 331 individuals. Anthropogenic noise is considered a threat for this population and can negatively impact communication. To characterize this population's vocal behavior, vocalizations were measured and classified into three categories: whistles (n = 1264, 77%), pulsed calls (n = 354, 22%), and combined calls (n = 15, 1%), resulting in 41 call types. Two quantitative analyses were conducted to compare with the manual classification. A classification and regression tree and Random Forest had a 95% and 85% agreement with the manual classification, respectively. The most common call types per category were then used to investigate masking by commercial ship noise. Results indicate that these call types were partially masked by distant ship noise and completely masked by close ship noise in the frequency range of 0-12 kHz. Understanding vocal behavior and the effects of masking in Cook Inlet belugas provides important information supporting the management of this endangered population.

Beluga whale (Delphinapterus leucas) acoustic foraging behavior and applications for long term monitoring.

Cook Inlet, Alaska, is home to an endangered and declining population of 279 belugas (Delphinapterus leucas). Recovery efforts highlight a paucity of basic ecological knowledge, impeding the correct assessment of threats and the development of recovery actions. In particular, information on diet and foraging habitat is very limited for this population. Passive acoustic monitoring has proven to be an efficient approach to monitor beluga distribution and seasonal occurrence. Identifying acoustic foraging behavior could help address the current gap in information on diet and foraging habitat. To address this conservation challenge, eight belugas from a comparative, healthy population in Bristol Bay, Alaska, were instrumented with a multi-sensor tag (DTAG), a satellite tag, and a stomach temperature transmitter in August 2014 and May 2016. DTAG deployments provided 129.6 hours of data including foraging and social behavioral states. A total of 68 echolocation click trains ending in terminal buzzes were identified during successful prey chasing and capture, as well as during social interactions. Of these, 37 click trains were successfully processed to measure inter-click intervals (ICI) and ICI trend in their buzzing section. Terminal buzzes with short ICI (minimum ICI <8.98 ms) and consistently decreasing ICI trend (ICI increment range <1.49 ms) were exclusively associated with feeding behavior. This dual metric was applied to acoustic data from one acoustic mooring within the Cook Inlet beluga critical habitat as an example of the application of detecting feeding in long-term passive acoustic monitoring data. This approach allowed description of the relationship between beluga presence, feeding occurrence, and the timing of spawning runs by different species of anadromous fish. Results reflected a clear preference for the Susitna River delta during eulachon (Thaleichthys pacificus), Chinook (Oncorhynchus tshawytscha), pink (Oncorhynchus gorbuscha), and coho (Oncorhynchus kisutch) salmon spawning run periods, with increased feeding occurrence at the peak of the Chinook and pink salmon runs.

Audiogram of a Cook Inlet beluga whale (Delphinapterus leucas).

Noise is a stressor to wildlife, yet the precise sound sensitivity of individuals and populations is often unknown or unmeasured. Cook Inlet, Alaska belugas (CIBs) are a critically endangered and declining marine mammal population. Anthropogenic noise is a primary threat to these animals. Auditory evoked potentials were used to measure the hearing of a wild, stranded CIB as part of its rehabilitation assessment. The beluga showed broadband (4-128 kHz) and sensitive hearing (<80 dB) for a wide-range of frequencies (16-80 kHz), reflective of a healthy odontocete auditory system. Data were similar to healthy, adult belugas from the comparative Bristol Bay population (the only other published data set of healthy, wild marine mammal hearing). Repeated October and December 2017 measurements were similar, showing continued auditory health of the animal throughout the rehabilitation period. Hearing data were compared to pile-driving and container-ship noise measurements made in Cook Inlet, two sources of concern, suggesting masking is likely at ecologically relevant distances. These data provide the first empirical hearing data for a CIB allowing for estimations of sound-sensitivity in this population. The beluga's sensitive hearing and likelihood of masking show noise is a clear concern for this population struggling to recover.

Beluga whale acoustic signal classification using deep learning neural network models.

Over a decade after the Cook Inlet beluga (Delphinapterus leucas) was listed as endangered in 2008, the population has shown no sign of recovery. Lack of ecological knowledge limits the understanding of, and ability to manage, potential threats impeding recovery of this declining population. National Oceanic and Atmospheric Administration Fisheries, in partnership with the Alaska Department of Fish and Game, initiated a passive acoustics monitoring program in 2017 to investigate beluga seasonal occurrence by deploying a series of passive acoustic moorings. Data have been processed with semi-automated tonal detectors followed by time intensive manual validation. To reduce this labor intensive and time-consuming process, in addition to increasing the accuracy of classification results, the authors constructed an ensembled deep learning convolutional neural network model to classify beluga detections as true or false. Using a 0.5 threshold, the final model achieves 96.57% precision and 92.26% recall on testing dataset. This methodology proves to be successful at classifying beluga signals, and the framework can be easily generalized to other acoustic classification problems.

Variation in hearing within a wild population of beluga whales (Delphinapterus leucas).

Documenting hearing abilities is vital to understanding a species' acoustic ecology and for predicting the impacts of increasing anthropogenic noise. Cetaceans use sound for essential biological functions such as foraging, navigation and communication; hearing is considered to be their primary sensory modality. Yet, we know little regarding the hearing of most, if not all, cetacean populations, which limits our understanding of their sensory ecology, population level variability and the potential impacts of increasing anthropogenic noise. We obtained audiograms (5.6-150 kHz) of 26 wild beluga whales to measure hearing thresholds during capture-release events in Bristol Bay, AK, USA, using auditory evoked potential methods. The goal was to establish the baseline population audiogram, incidences of hearing loss and general variability in wild beluga whales. In general, belugas showed sensitive hearing with low thresholds (<80 dB) from 16 to 100 kHz, and most individuals (76%) responded to at least 120 kHz. Despite belugas often showing sensitive hearing, thresholds were usually above or approached the low ambient noise levels measured in the area, suggesting that a quiet environment may be associated with hearing sensitivity and that hearing thresholds in the most sensitive animals may have been masked. Although this is just one wild population, the success of the method suggests that it should be applied to other populations and species to better assess potential differences. Bristol Bay beluga audiograms showed substantial (30-70 dB) variation among individuals; this variation increased at higher frequencies. Differences among individual belugas reflect that testing multiple individuals of a population is necessary to best describe maximum sensitivity and population variance. The results of this study quadruple the number of individual beluga whales for which audiograms have been conducted and provide the first auditory data for a population of healthy wild odontocetes.

Dual instrument passive acoustic monitoring of belugas in Cook Inlet, Alaska.

As part of a long-term research program, Cook Inlet beluga (Delphinapterus leucas) presence was acoustically monitored with two types of acoustic sensors utilized in tandem in moorings deployed year-round: an ecological acoustic recorder (EAR) and a cetacean and porpoise detector (C-POD). The EAR was used primarily to record the calls, whistles, and buzzes produced by belugas and killer whales (Orcinus orca). The C-POD was used to log and classify echolocation clicks from belugas, killer whales, and porpoises. This paper describes mooring packages that maximized the chances of successful long-term data collection in the particularly challenging Cook Inlet environment, and presents an analytical comparison of odontocete detections obtained by the collocated EAR and C-POD instruments from two mooring locations in the upper inlet. Results from this study illustrate a significant improvement in detecting beluga and killer whale presence when the different acoustic signals detected by EARs and C-PODs are considered together. Further, results from concurrent porpoise detections indicating prey competition and feeding interference with beluga, and porpoise displacement due to ice formation are described.

Review of the Effects of Offshore Seismic Surveys in Cetaceans: Are Mass Strandings a Possibility?

Displacement of cetaceans is commonly reported during offshore seismic surveys. Speculation concerning possible links between seismic survey noise and cetacean strandings is available for a dozen events but without convincing causal evidence. This lack of evidence should not be considered conclusive but rather as reflecting the absence of a comprehensive analysis of the circumstances. Current mitigation guidelines are inadequate for long-range effects such as displacements and the potential for strandings. This review presents the available information for ten documented strandings that were possibly linked to seismic surveys and recommends initial measures and actions to further evaluate this potential link.

The European Marine Strategy: Noise Monitoring in European Marine Waters from 2014.

The European Marine Strategy Framework Directive requires European member states to develop strategies for their marine waters leading to programs of measures that achieve or maintain good environmental status (GES) in all European seas by 2020. An essential step toward reaching GES is the establishment of monitoring programs, enabling the state of marine waters to be assessed on a regular basis. A register for impulsive noise-generating activities would enable assessment of their cumulative impacts on wide temporal and spatial scales; monitoring of ambient noise would provide essential insight into current levels and any trend in European waters.

Measuring Hearing in Wild Beluga Whales.

We measured the hearing abilities of seven wild beluga whales (Delphinapterus leucas) during a collection-and-release experiment in Bristol Bay, AK. Here we summarize the methods and initial data from one animal and discuss the implications of this experiment. Audiograms were collected from 4 to 150 kHz. The animal with the lowest threshold heard best at 80 kHz and demonstrated overall good hearing from 22 to 110 kHz. The robustness of the methodology and data suggest that the auditory evoked potential audiograms can be incorporated into future collection-and-release health assessments. Such methods may provide high-quality results for multiple animals, facilitating population-level audiograms and hearing measures in new species.

Beluga whale (Delphinapterus leucas) vocalizations and call classification from the eastern Beaufort Sea population.

Beluga whales, Delphinapterus leucas, have a graded call system; call types exist on a continuum making classification challenging. A description of vocalizations from the eastern Beaufort Sea beluga population during its spring migration are presented here, using both a non-parametric classification tree analysis (CART), and a Random Forest analysis. Twelve frequency and duration measurements were made on 1019 calls recorded over 14 days off Icy Cape, Alaska, resulting in 34 identifiable call types with 83% agreement in classification for both CART and Random Forest analyses. This high level of agreement in classification, with an initial subjective classification of calls into 36 categories, demonstrates that the methods applied here provide a quantitative analysis of a graded call dataset. Further, as calls cannot be attributed to individuals using single sensor passive acoustic monitoring efforts, these methods provide a comprehensive analysis of data where the influence of pseudo-replication of calls from individuals is unknown. This study is the first to describe the vocal repertoire of a beluga population using a robust and repeatable methodology. A baseline eastern Beaufort Sea beluga population repertoire is presented here, against which the call repertoire of other seasonally sympatric Alaskan beluga populations can be compared.

Baseline hearing abilities and variability in wild beluga whales (Delphinapterus leucas).

While hearing is the primary sensory modality for odontocetes, there are few data addressing variation within a natural population. This work describes the hearing ranges (4-150 kHz) and sensitivities of seven apparently healthy, wild beluga whales (Delphinapterus leucas) during a population health assessment project that captured and released belugas in Bristol Bay, Alaska. The baseline hearing abilities and subsequent variations were addressed. Hearing was measured using auditory evoked potentials (AEPs). All audiograms showed a typical cetacean U-shape; substantial variation (>30 dB) was found between most and least sensitive thresholds. All animals heard well, up to at least 128 kHz. Two heard up to 150 kHz. Lowest auditory thresholds (35-45 dB) were identified in the range 45-80 kHz. Greatest differences in hearing abilities occurred at both the high end of the auditory range and at frequencies of maximum sensitivity. In general, wild beluga hearing was quite sensitive. Hearing abilities were similar to those of belugas measured in zoological settings, reinforcing the comparative importance of both settings. The relative degree of variability across the wild belugas suggests that audiograms from multiple individuals are needed to properly describe the maximum sensitivity and population variance for odontocetes. Hearing measures were easily incorporated into field-based settings. This detailed examination of hearing abilities in wild Bristol Bay belugas provides a basis for a better understanding of the potential impact of anthropogenic noise on a noise-sensitive species. Such information may help design noise-limiting mitigation measures that could be applied to areas heavily influenced and inhabited by endangered belugas.

Seasonal migrations of North Atlantic minke whales: novel insights from large-scale passive acoustic monitoring networks.

BACKGROUND: Little is known about migration patterns and seasonal distribution away from coastal summer feeding habitats of many pelagic baleen whales. Recently, large-scale passive acoustic monitoring networks have become available to explore migration patterns and identify critical habitats of these species. North Atlantic minke whales (Balaenoptera acutorostrata) perform seasonal migrations between high latitude summer feeding and low latitude winter breeding grounds. While the distribution and abundance of the species has been studied across their summer range, data on migration and winter habitat are virtually missing. Acoustic recordings, from 16 different sites from across the North Atlantic, were analyzed to examine the seasonal and geographic variation in minke whale pulse train occurrence, infer information about migration routes and timing, and to identify possible winter habitats. RESULTS: Acoustic detections show that minke whales leave their winter grounds south of 30° N from March through early April. On their southward migration in autumn, minke whales leave waters north of 40° N from mid-October through early November. In the western North Atlantic spring migrants appear to track the warmer waters of the Gulf Stream along the continental shelf, while whales travel farther offshore in autumn. Abundant detections were found off the southeastern US and the Caribbean during winter. Minke whale pulse trains showed evidence of geographic variation, with longer pulse trains recorded south of 40° N. Very few pulse trains were recorded during summer in any of the datasets. CONCLUSION: This study highlights the feasibility of using acoustic monitoring networks to explore migration patterns of pelagic marine mammals. Results confirm the presence of minke whales off the southeastern US and the Caribbean during winter months. The absence of pulse train detections during summer suggests either that minke whales switch their vocal behaviour at this time of year, are absent from available recording sites or that variation in signal structure influenced automated detection. Alternatively, if pulse trains are produced in a reproductive context by males, these data may indicate their absence from the selected recording sites. Evidence of geographic variation in pulse train duration suggests different behavioural functions or use of these calls at different latitudes.

Passive acoustic monitoring of Cook Inlet beluga whales (Delphinapterus leucas).

The endangered beluga whale (Delphinapterus leucas) population in Cook Inlet, AK faces threats from a variety of anthropogenic factors, including coastal development, oil and gas exploration, vessel traffic, and military activities. To address existing gaps in understanding about the occurrence of belugas in Cook Inlet, a project was developed to use passive acoustic monitoring to document the year-round distribution of belugas, as well as killer whales (Orcinus orca), which prey on belugas. Beginning in June 2009, ten moorings were deployed throughout the Inlet and refurbished every two to eight months. Despite challenging conditions consisting of strong tidal currents carrying debris and seasonal ice cover, 83% of mooring deployments were successfully recovered. Noise from water flow, vessel traffic, and/or industrial activities was present at several sites, potentially masking some signals. However, belugas were successfully detected at multiple locations. Detections were relatively common in the upper inlet and less common or absent at middle and lower inlet locations. Killer whale signals were also recorded. Some seasonal variability in the occurrence of both belugas and killer whales was evident.

Evidence that ship noise increases stress in right whales.

Baleen whales (Mysticeti) communicate using low-frequency acoustic signals. These long-wavelength sounds can be detected over hundreds of kilometres, potentially allowing contact over large distances. Low-frequency noise from large ships (20-200 Hz) overlaps acoustic signals used by baleen whales, and increased levels of underwater noise have been documented in areas with high shipping traffic. Reported responses of whales to increased noise include: habitat displacement, behavioural changes and alterations in the intensity, frequency and intervals of calls. However, it has been unclear whether exposure to noise results in physiological responses that may lead to significant consequences for individuals or populations. Here, we show that reduced ship traffic in the Bay of Fundy, Canada, following the events of 11 September 2001, resulted in a 6 dB decrease in underwater noise with a significant reduction below 150 Hz. This noise reduction was associated with decreased baseline levels of stress-related faecal hormone metabolites (glucocorticoids) in North Atlantic right whales (Eubalaena glacialis). This is the first evidence that exposure to low-frequency ship noise may be associated with chronic stress in whales, and has implications for all baleen whales in heavy ship traffic areas, and for recovery of this endangered right whale population.

The beluga whale produces two pulses to form its sonar signal.

Odontocete cetaceans use biosonar clicks to acoustically probe their aquatic environment with an aptitude unmatched by man-made sonar. A cornerstone of this ability is their use of short, broadband pulses produced in the region of the upper nasal passages. Here we provide empirical evidence that a beluga whale (Delphinapterus leucas) uses two signal generators simultaneously when echolocating. We show that the pulses of the two generators are combined as they are transmitted through the melon to produce a single echolocation click emitted from the front of the animal. Generating two pulses probably offers the beluga the ability to control the energy and frequency distribution of the emitted click and may allow it to acoustically steer its echolocation beam.