Vulnerability to climate change of United States marine mammal stocks in the western North Atlantic, Gulf of Mexico, and Caribbean.

Climate change and climate variability are affecting marine mammal species and these impacts are projected to continue in the coming decades. Vulnerability assessments provide a framework for evaluating climate impacts over a broad range of species using currently available information. We conducted a trait-based climate vulnerability assessment using expert elicitation for 108 marine mammal stocks and stock groups in the western North Atlantic, Gulf of Mexico, and Caribbean Sea. Our approach combined the exposure (projected change in environmental conditions) and sensitivity (ability to tolerate and adapt to changing conditions) of marine mammal stocks to estimate vulnerability to climate change, and categorize stocks with a vulnerability index. The climate vulnerability score was very high for 44% (n = 47) of these stocks, high for 29% (n = 31), moderate for 20% (n = 22), and low for 7% (n = 8). The majority of stocks (n = 78; 72%) scored very high exposure, whereas 24% (n = 26) scored high, and 4% (n = 4) scored moderate. The sensitivity score was very high for 33% (n = 36) of these stocks, high for 18% (n = 19), moderate for 34% (n = 37), and low for 15% (n = 16). Vulnerability results were summarized for stocks in five taxonomic groups: pinnipeds (n = 4; 25% high, 75% moderate), mysticetes (n = 7; 29% very high, 57% high, 14% moderate), ziphiids (n = 8; 13% very high, 50% high, 38% moderate), delphinids (n = 84; 52% very high, 23% high, 15% moderate, 10% low), and other odontocetes (n = 5; 60% high, 40% moderate). Factors including temperature, ocean pH, and dissolved oxygen were the primary drivers of high climate exposure, with effects mediated through prey and habitat parameters. We quantified sources of uncertainty by bootstrapping vulnerability scores, conducting leave-one-out analyses of individual attributes and individual scorers, and through scoring data quality for each attribute. These results provide information for researchers, managers, and the public on marine mammal responses to climate change to enhance the development of more effective marine mammal management, restoration, and conservation activities that address current and future environmental variation and biological responses due to climate change.

Kinematics and energetics of foraging behavior in Rice's whales of the Gulf of Mexico.

Rorqual foraging behavior varies with species, prey type and foraging conditions, and can be a determining factor for their fitness. Little is known about the foraging ecology of Rice's whales (Balaenoptera ricei), an endangered species with a population of fewer than 100 individuals. Suction cup tags were attached to two Rice's whales to collect information on their diving kinematics and foraging behavior. The tagged whales primarily exhibited lunge-feeding near the sea bottom and to a lesser extent in the water-column and at the sea surface. During 6-10 min foraging dives, the whales typically circled their prey before executing one or two feeding lunges. Longer duration dives and dives with more feeding-lunges were followed by an increase in their breathing rate. The median lunge rate of one lunge per dive of both animals was much lower than expected based on comparative research on other lunge-feeding baleen whales, and may be associated with foraging on fish instead of krill or may be an indication of different foraging conditions. Both animals spent extended periods of the night near the sea surface, increasing the risk for ship strike. Furthermore, their circling before lunging may increase the risk for entanglement in bottom-longline fishing gear. Overall, these data show that Rice's whale foraging behavior differs from other lunge feeding rorqual species and may be a significant factor in shaping our understanding of their foraging ecology. Efforts to mitigate threats to Rice's whales will benefit from improved understanding of patterns in their habitat use and fine-scale ecology.

Modeling protected species distributions and habitats to inform siting and management of pioneering ocean industries: A case study for Gulf of Mexico aquaculture.

Marine Spatial Planning (MSP) provides a process that uses spatial data and models to evaluate environmental, social, economic, cultural, and management trade-offs when siting (i.e., strategically locating) ocean industries. Aquaculture is the fastest-growing food sector in the world. The United States (U.S.) has substantial opportunity for offshore aquaculture development given the size of its exclusive economic zone, habitat diversity, and variety of candidate species for cultivation. However, promising aquaculture areas overlap many protected species habitats. Aquaculture siting surveys, construction, operations, and decommissioning can alter protected species habitat and behavior. Additionally, aquaculture-associated vessel activity, underwater noise, and physical interactions between protected species and farms can increase the risk of injury and mortality. In 2020, the U.S. Gulf of Mexico was identified as one of the first regions to be evaluated for offshore aquaculture opportunities as directed by a Presidential Executive Order. We developed a transparent and repeatable method to identify aquaculture opportunity areas (AOAs) with the least conflict with protected species. First, we developed a generalized scoring approach for protected species that captures their vulnerability to adverse effects from anthropogenic activities using conservation status and demographic information. Next, we applied this approach to data layers for eight species listed under the Endangered Species Act, including five species of sea turtles, Rice's whale, smalltooth sawfish, and giant manta ray. Next, we evaluated four methods for mathematically combining scores (i.e., Arithmetic mean, Geometric mean, Product, Lowest Scoring layer) to generate a combined protected species data layer. The Product approach provided the most logical ordering of, and the greatest contrast in, site suitability scores. Finally, we integrated the combined protected species data layer into a multi-criteria decision-making modeling framework for MSP. This process identified AOAs with reduced potential for protected species conflict. These modeling methods are transferable to other regions, to other sensitive or protected species, and for spatial planning for other ocean-uses.

Acoustic localization, validation, and characterization of Rice's whale calls.

The recently named Rice's whale in the Gulf of Mexico is one of the most endangered whales in the world, and improved knowledge of spatiotemporal occurrence patterns is needed to support their recovery and conservation. Passive acoustic monitoring methods for determining spatiotemporal occurrence patterns require identifying the species' call repertoire. Rice's whale call repertoire remains unvalidated though several potential call types have been identified. This study uses sonobuoys and passive acoustic tagging to validate the source of potential call types and to characterize Rice's whale calls. During concurrent visual and acoustic surveys, acoustic-directed approaches were conducted to obtain visual verifications of sources of localized sounds. Of 28 acoustic-directed approaches, 79% led to sightings of balaenopterid whales, of which 10 could be positively identified to species as Rice's whales. Long-moan calls, downsweep sequences, and tonal-sequences are attributed to Rice's whales based on these matches, while anthropogenic sources are ruled out. A potential new call type, the low-frequency downsweep sequence, is characterized from tagged Rice's whale recordings. The validation and characterization of the Rice's whale call repertoire provides foundational information needed to use passive acoustic monitoring for better understanding and conservation of these critically endangered whales.

Cetacean distribution models based on visual and passive acoustic data.

Distribution models are needed to understand spatiotemporal patterns in cetacean occurrence and to mitigate anthropogenic impacts. Shipboard line-transect visual surveys are the standard method for estimating abundance and describing the distributions of cetacean populations. Ship-board surveys provide high spatial resolution but lack temporal resolution and seasonal coverage. Stationary passive acoustic monitoring (PAM) employs acoustic sensors to sample point locations nearly continuously, providing high temporal resolution in local habitats across days, seasons and years. To evaluate whether cross-platform data synthesis can improve distribution predictions, models were developed for Cuvier's beaked whales, sperm whales, and Risso's dolphins in the oceanic Gulf of Mexico using two different methods: generalized additive models and neural networks. Neural networks were able to learn unspecified interactions between drivers. Models that incorporated PAM datasets out-performed models trained on visual data alone, and joint models performed best in two out of three cases. The modeling results suggest that, when taken together, multiple species distribution models using a variety of data types may support conservation and management of Gulf of Mexico cetacean populations by improving the understanding of temporal and spatial species distribution trends.

Exploring movement patterns and changing distributions of baleen whales in the western North Atlantic using a decade of passive acoustic data.

Six baleen whale species are found in the temperate western North Atlantic Ocean, with limited information existing on the distribution and movement patterns for most. There is mounting evidence of distributional shifts in many species, including marine mammals, likely because of climate-driven changes in ocean temperature and circulation. Previous acoustic studies examined the occurrence of minke (Balaenoptera acutorostrata) and North Atlantic right whales (NARW; Eubalaena glacialis). This study assesses the acoustic presence of humpback (Megaptera novaeangliae), sei (B. borealis), fin (B. physalus), and blue whales (B. musculus) over a decade, based on daily detections of their vocalizations. Data collected from 2004 to 2014 on 281 bottom-mounted recorders, totaling 35,033 days, were processed using automated detection software and screened for each species' presence. A published study on NARW acoustics revealed significant changes in occurrence patterns between the periods of 2004-2010 and 2011-2014; therefore, these same time periods were examined here. All four species were present from the Southeast United States to Greenland; humpback whales were also present in the Caribbean. All species occurred throughout all regions in the winter, suggesting that baleen whales are widely distributed during these months. Each of the species showed significant changes in acoustic occurrence after 2010. Similar to NARWs, sei whales had higher acoustic occurrence in mid-Atlantic regions after 2010. Fin, blue, and sei whales were more frequently detected in the northern latitudes of the study area after 2010. Despite this general northward shift, all four species were detected less on the Scotian Shelf area after 2010, matching documented shifts in prey availability in this region. A decade of acoustic observations have shown important distributional changes over the range of baleen whales, mirroring known climatic shifts and identifying new habitats that will require further protection from anthropogenic threats like fixed fishing gear, shipping, and noise pollution.

Displaying bioacoustic directional information from sonobuoys using "azigrams".

The AN/SSQ-53 Directional Frequency Analysis and Recording (DIFAR) sonobuoy is an expendable device that can derive acoustic particle velocity along two orthogonal horizontal axes, along with acoustic pressure. This information enables computation of azimuths of low-frequency acoustic sources from a single compact sensor. The standard approach for estimating azimuth from these sensors is by conventional beamforming (i.e., adding weighted time series), but the resulting "cardioid" beampattern is imprecise, computationally expensive, and vulnerable to directional noise contamination for weak signals. Demonstrated here is an alternative multiplicative processing scheme that computes the "active intensity" of an acoustic signal to obtain the dominant directionality of a noise field as a function of time and frequency. This information is conveniently displayed as an "azigram," which is analogous to a spectrogram, but uses color to indicate azimuth instead of intensity. Data from several locations demonstrate this approach, which can be computed without demultiplexing the raw signal. Azigrams have been used to help diagnose sonobuoy issues, improve detectability, and estimate bearings of low signal-to-noise ratio signals. Azigrams may also enhance the detection and potential classification of signals embedded in directional noise fields.

Automated classification of dolphin echolocation click types from the Gulf of Mexico.

Delphinids produce large numbers of short duration, broadband echolocation clicks which may be useful for species classification in passive acoustic monitoring efforts. A challenge in echolocation click classification is to overcome the many sources of variability to recognize underlying patterns across many detections. An automated unsupervised network-based classification method was developed to simulate the approach a human analyst uses when categorizing click types: Clusters of similar clicks were identified by incorporating multiple click characteristics (spectral shape and inter-click interval distributions) to distinguish within-type from between-type variation, and identify distinct, persistent click types. Once click types were established, an algorithm for classifying novel detections using existing clusters was tested. The automated classification method was applied to a dataset of 52 million clicks detected across five monitoring sites over two years in the Gulf of Mexico (GOM). Seven distinct click types were identified, one of which is known to be associated with an acoustically identifiable delphinid (Risso's dolphin) and six of which are not yet identified. All types occurred at multiple monitoring locations, but the relative occurrence of types varied, particularly between continental shelf and slope locations. Automatically-identified click types from autonomous seafloor recorders without verifiable species identification were compared with clicks detected on sea-surface towed hydrophone arrays in the presence of visually identified delphinid species. These comparisons suggest potential species identities for the animals producing some echolocation click types. The network-based classification method presented here is effective for rapid, unsupervised delphinid click classification across large datasets in which the click types may not be known a priori.

Long-term passive acoustic recordings track the changing distribution of North Atlantic right whales (Eubalaena glacialis) from 2004 to 2014.

Given new distribution patterns of the endangered North Atlantic right whale (NARW; Eubalaena glacialis) population in recent years, an improved understanding of spatio-temporal movements are imperative for the conservation of this species. While so far visual data have provided most information on NARW movements, passive acoustic monitoring (PAM) was used in this study in order to better capture year-round NARW presence. This project used PAM data from 2004 to 2014 collected by 19 organizations throughout the western North Atlantic Ocean. Overall, data from 324 recorders (35,600 days) were processed and analyzed using a classification and detection system. Results highlight almost year-round habitat use of the western North Atlantic Ocean, with a decrease in detections in waters off Cape Hatteras, North Carolina in summer and fall. Data collected post 2010 showed an increased NARW presence in the mid-Atlantic region and a simultaneous decrease in the northern Gulf of Maine. In addition, NARWs were widely distributed across most regions throughout winter months. This study demonstrates that a large-scale analysis of PAM data provides significant value to understanding and tracking shifts in large whale movements over long time scales.

Geographic variation in Risso's dolphin echolocation click spectra.

Discrimination of bioacoustic signals to the species or population level is critical for using passive acoustic monitoring to study cetacean ecology. Risso's dolphins off southern California have distinctive peaks and notches in their echolocation clicks, but it was unknown whether Risso's dolphins from other geographic areas have similarly distinctive click spectra and whether populations are acoustically distinct. This study investigates using clicks for species and population identification by characterizing the spectral structure of Risso's dolphin echolocation clicks recorded over wide-ranging geographic regions including the U.S. waters of the North Atlantic Ocean, Gulf of Mexico, and North Pacific Ocean; and international waters of the Eastern Tropical Pacific. All recordings with Risso's dolphin clicks exhibited the spectral peak and notch pattern described off southern California, indicating the presence of peak banding patterns is useful for species discrimination. Geographic regions were a significant explanatory factor for variability in the frequencies of click spectral peaks, with relatively higher frequency peaks and notches found off Hawaii compared to California waters and off the southeast U.S. compared to the Gulf of Mexico. In the North Atlantic Ocean, a latitudinal cline in frequencies was evident. Potential causes of acoustic variation within and among acoustic encounters are evaluated.

Automated extraction of odontocete whistle contours.

Many odontocetes produce frequency modulated tonal calls known as whistles. The ability to automatically determine time × frequency tracks corresponding to these vocalizations has numerous applications including species description, identification, and density estimation. This work develops and compares two algorithms on a common corpus of nearly one hour of data collected in the Southern California Bight and at Palmyra Atoll. The corpus contains over 3000 whistles from bottlenose dolphins, long- and short-beaked common dolphins, spinner dolphins, and melon-headed whales that have been annotated by a human, and released to the Moby Sound archive. Both algorithms use a common signal processing front end to determine time × frequency peaks from a spectrogram. In the first method, a particle filter performs Bayesian filtering, estimating the contour from the noisy spectral peaks. The second method uses an adaptive polynomial prediction to connect peaks into a graph, merging graphs when they cross. Whistle contours are extracted from graphs using information from both sides of crossings. The particle filter was able to retrieve 71.5% (recall) of the human annotated tonals with 60.8% of the detections being valid (precision). The graph algorithm's recall rate was 80.0% with a precision of 76.9%.

Classification of echolocation clicks from odontocetes in the Southern California Bight.

This study presents a system for classifying echolocation clicks of six species of odontocetes in the Southern California Bight: Visually confirmed bottlenose dolphins, short- and long-beaked common dolphins, Pacific white-sided dolphins, Risso's dolphins, and presumed Cuvier's beaked whales. Echolocation clicks are represented by cepstral feature vectors that are classified by Gaussian mixture models. A randomized cross-validation experiment is designed to provide conditions similar to those found in a field-deployed system. To prevent matched conditions from inappropriately lowering the error rate, echolocation clicks associated with a single sighting are never split across the training and test data. Sightings are randomly permuted before assignment to folds in the experiment. This allows different combinations of the training and test data to be used while keeping data from each sighting entirely in the training or test set. The system achieves a mean error rate of 22% across 100 randomized three-fold cross-validation experiments. Four of the six species had mean error rates lower than the overall mean, with the presumed Cuvier's beaked whale clicks showing the best performance (<2% error rate). Long-beaked common and bottlenose dolphins proved the most difficult to classify, with mean error rates of 53% and 68%, respectively.

Spatial and temporal patterns of Risso's dolphin echolocation in the Southern California Bight.

Geographical and temporal trends in echolocation clicking activity can lead to insights into the foraging and migratory behaviors of pelagic dolphins. Using autonomous acoustic recording packages, the geographical, diel, and seasonal patterns of Risso's dolphin (Grampus griseus) echolocation click activity are described for six locations in the Southern California Bight between 2005 and 2007. Risso's dolphin echolocation click bouts are identified based on their unique spectral characteristics. Click bouts were identified on 739 of 1959 recording days at all 6 sites, with the majority occurring at nearshore sites. A significant diel pattern is evident in which both hourly occurrences of click bouts and click rates are higher at night than during the day. At all nearshore sites, Risso's dolphin clicks were identified year-round, with the highest daily occurrence at the southern end of Santa Catalina Island. Seasonal and interannual variabilities in occurrence were high across sites with peak occurrence in autumn of most years at most sites. These results suggest that Risso's dolphins forage at night and that the southern end of Santa Catalina Island represents an important habitat for Risso's dolphins throughout the year.

Classification of Risso's and Pacific white-sided dolphins using spectral properties of echolocation clicks.

The spectral and temporal properties of echolocation clicks and the use of clicks for species classification are investigated for five species of free-ranging dolphins found offshore of southern California: short-beaked common (Delphinus delphis), long-beaked common (D. capensis), Risso's (Grampus griseus), Pacific white-sided (Lagenorhynchus obliquidens), and bottlenose (Tursiops truncatus) dolphins. Spectral properties are compared among the five species and unique spectral peak and notch patterns are described for two species. The spectral peak mean values from Pacific white-sided dolphin clicks are 22.2, 26.6, 33.7, and 37.3 kHz and from Risso's dolphins are 22.4, 25.5, 30.5, and 38.8 kHz. The spectral notch mean values from Pacific white-sided dolphin clicks are 19.0, 24.5, and 29.7 kHz and from Risso's dolphins are 19.6, 27.7, and 35.9 kHz. Analysis of variance analyses indicate that spectral peaks and notches within the frequency band 24-35 kHz are distinct between the two species and exhibit low variation within each species. Post hoc tests divide Pacific white-sided dolphin recordings into two distinct subsets containing different click types, which are hypothesized to represent the different populations that occur within the region. Bottlenose and common dolphin clicks do not show consistent patterns of spectral peaks or notches within the frequency band examined (1-100 kHz).

Anatomic geometry of sound transmission and reception in Cuvier's beaked whale (Ziphius cavirostris).

This study uses remote imaging technology to quantify, compare, and contrast the cephalic anatomy between a neonate female and a young adult male Cuvier's beaked whale. Primary results reveal details of anatomic geometry with implications for acoustic function and diving. Specifically, we describe the juxtaposition of the large pterygoid sinuses, a fibrous venous plexus, and a lipid-rich pathway that connects the acoustic environment to the bony ear complex. We surmise that the large pterygoid air sinuses are essential adaptations for maintaining acoustic isolation and auditory acuity of the ears at depth. In the adult male, an acoustic waveguide lined with pachyosteosclerotic bones is apparently part of a novel transmission pathway for outgoing biosonar signals. Substitution of dense tissue boundaries where we normally find air sacs in delphinoids appears to be a recurring theme in deep-diving beaked whales and sperm whales. The anatomic configuration of the adult male Ziphius forehead resembles an upside-down sperm whale nose and may be its functional equivalent, but the homologous relationships between forehead structures are equivocal.

Gaussian mixture model classification of odontocetes in the Southern California Bight and the Gulf of California.

A method for the automatic classification of free-ranging delphinid vocalizations is presented. The vocalizations of short-beaked and long-beaked common (Delphinus delphis and Delphinus capensis), Pacific white-sided (Lagenorhynchus obliquidens), and bottlenose (Tursiops truncatus) dolphins were recorded in a pelagic environment of the Southern California Bight and the Gulf of California over a period of 4 years. Cepstral feature vectors are extracted from call data which contain simultaneous overlapping whistles, burst-pulses, and clicks from a single species. These features are grouped into multisecond segments. A portion of the data is used to train Gaussian mixture models of varying orders for each species. The remaining call data are used to test the performance of the models. Species are predicted based upon probabilistic measures of model similarity with test segment groups having durations between 1 and 25 s. For this data set, 256 mixture Gaussian mixture models and segments of at least 10 s of call data resulted in the best classification results. The classifier predicts the species of groups with 67%-75% accuracy depending upon the partitioning of the training and test data.

Cuvier's beaked whale (Ziphius cavirostris) head tissues: physical properties and CT imaging.

Tissue physical properties from a Cuvier's beaked whale (Ziphius cavirostris) neonate head are reported and compared with computed tomography (CT) X-ray imaging. Physical properties measured include longitudinal sound velocity, density, elastic modulus and hysteresis. Tissues were classified by type as follows: mandibular acoustic fat, mandibular blubber, forehead acoustic fat (melon), forehead blubber, muscle and connective tissue. Results show that each class of tissues has unique, co-varying physical properties. The mandibular acoustic fats had minimal values for sound speed (1350+/-10.6 m s(-1)) and mass density (890+/-23 kg m(-3)). These values increased through mandibular blubber (1376+/-13 m s(-1), 919+/-13 kg m(-3)), melon (1382+/-23 m s(-1), 937+/-17 kg m(-3)), forehead blubber (1401+/-7.8 m s(-1), 935+/-25 kg m(-3)) and muscle (1517+/-46.8 m s(-1), 993+/-58 kg m(-3)). Connective tissue had the greatest mean sound speed and density (1628+/-48.7 m s(-1), 1087+/-41 kg m(-3)). The melon formed a low-density, low-sound-speed core, supporting its function as a sound focusing organ. Hounsfield unit (HU) values from CT X-ray imaging are correlated with density and sound speed values, allowing HU values to be used to predict these physical properties. Blubber and connective tissues have a higher elastic modulus than acoustic fats and melon, suggesting more collagen structure in blubber and connective tissues. Blubber tissue elastic modulus is nonlinear with varying stress, becoming more incompressible as stress is increased. These data provide important physical properties required to construct models of the sound generation and reception mechanisms in Ziphius cavirostris heads, as well as models of their interaction with anthropogenic sound.