Geographic distribution of the Cross Seamount beaked whale based on acoustic detections

Beaked whales produce frequency-modulated echolocation pulses that appear to be species-specific, allowing passive acoustic monitoring to play a role in understanding spatio-temporal patterns. The Cross Seamount beaked whale is known only from its unique echolocation signal (BWC) with no confirmed species identification. This beaked whale spans the Pacific Ocean from the Mariana Archipelago to Baja California, Mexico, south to the equator, but only as far north as latitude 29°N. Within these warm waters, 92% of BWC detections occurred at night, 6% during crepuscular periods, and only 2% during daylight hours. Detections of BWC signals on drifting recorders with a vertical hydrophone array at 150 m depth demonstrated that foraging often occurred shallow in the water column (<150 m). No other species of beaked whale to date has been documented foraging in waters this shallow. Given their nocturnal, shallow foraging dives, this species appears to prefer prey that may be available in the water column only during those hours. The foraging behavior of Cross Seamount beaked whales appears to be unique among all beaked whales, and these findings contribute additional ecological and acoustic information which can help guide future efforts to identify this cryptic whale.     

Deep-diving beaked whales dive together but forage apart

Echolocating animals that forage in social groups can potentially benefit from eavesdropping on other group members, cooperative foraging or social defence, but may also face problems of acoustic interference and intra-group competition for prey. Here, we investigate these potential trade-offs of sociality for extreme deep-diving Blainville′s and Cuvier''s beaked whales. These species perform highly synchronous group dives as a presumed predator-avoidance behaviour, but the benefits and costs of this on foraging have not been investigated. We show that group members could hear their companions for a median of at least 91% of the vocal foraging phase of their dives. This enables whales to coordinate their mean travel direction despite differing individual headings as they pursue prey on a minute-by-minute basis. While beaked whales coordinate their echolocation-based foraging periods tightly, individual click and buzz rates are both independent of the number of whales in the group. Thus, their foraging performance is not affected by intra-group competition or interference from group members, and they do not seem to capitalize directly on eavesdropping on the echoes produced by the echolocation clicks of their companions. We conclude that the close diving and vocal synchronization of beaked whale groups that quantitatively reduces predation risk has little impact on foraging performance.     

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.     

Seasonal acoustic behavior of blue whales (Balaenoptera musculus) in the Gulf of California,Mexico

Six years of passive acoustic monitoring data from the Gulf of California reveal seasonality and movements for the northeastern Pacific blue whales. Three sites were studied, one from the southern (Punta Pescadero) and two from the northern (Isla Tiburon and Canal de Ballenas) regions. A total of 4,953 h were analyzed, and 15,539 blue whale calls were detected, of which 2,723 (18%) were A calls, 11,249 were B calls (72%), and 1,567 were D calls (10%). A and B calls were produced both as song units (2,524) or AB singular calls (2,026). The high rate of songs and their seasonality suggest that the GC is a winter‐breeding ground. A shift from AB call predominance in winter, to D calls in spring and early summer, especially at the entrance of the GC, suggests the importance of this area for reproduction and foraging. Analysis of calling frequency suggests a clear movement of blue whales from the southern region (Punta Pescadero) to the northern regions (Canal de Ballenas and Isla Tiburon), with subsequent southern movement in March. The seasonality and mobility of blue whales in the Gulf of California, inferred from their calling, contributes to the ecological understanding of this population.     

The extracranial arterial system in the heads of beaked whales,with implications on diving physiology and pathogenesis

Beaked whales are medium‐sized toothed whales that inhabit depths beyond the continental shelf; thus beaked whale strandings are relatively infrequent compared to those of other cetaceans. Beaked whales have been catapulted into the spotlight by their tendency to strand in association with naval sonar deployment. Studies have shown the presence of gas and fat emboli within the tissues and analysis of gas emboli is suggestive of nitrogen as the primary component. These findings are consistent with human decompression sickness (DCS) previously not thought possible in cetaceans. Because, tissue loading with nitrogen gas is paramount for the manifestation of DCS and nitrogen loading depends largely on the vascular perfusion of the tissues, we examined the anatomy of the extracranial arterial system using stranded carcasses of 16 beaked whales from five different species. Anatomic regions containing lipid and/or air spaces were prioritized as potential locations of nitrogen gas absorption due to the known solubility of nitrogen in adipose tissue and the nitrogen content of air, respectively. Attention was focused on the acoustic fat bodies and accessory sinus system on the ventral head. We found much of the arterial system of the head to contain arteries homologous to those found in domestic mammals. Robust arterial associations with lipid depots and air spaces occurred within the acoustic fat bodies of the lower jaw and pterygoid air sacs of the ventral head, respectively. Both regions contained extensive trabecular geometry with small arteries investing the trabeculae. Our findings suggest the presence of considerable surface area between the arterial system, and the intramandibular fat bodies and pterygoid air sacs. Our observations may provide support for the hypothesis that these structures play an important role in the exchange of nitrogen gas during diving. J. Morphol. 277:5–33, 2016. © 2015 Wiley Periodicals, Inc.     

Geographic differences in Blainville's beaked whale (Mesoplodon densirostris) echolocation clicks

Aim

Understanding cetacean species' distributions and population structure over space and time is necessary for effective conservation and management. Geographic differences in acoustic signals may provide a line of evidence for population-level discrimination in some cetacean species. We use acoustic recordings collected over broad spatial and temporal scales to investigate whether global variability in echolocation click peak frequency could elucidate population structure in Blainville's beaked whale (Mesoplodon densirostris), a cryptic species well-studied acoustically.

Location

North Pacific, Western North Atlantic and Gulf of Mexico.

Time period

2004–2021.

Major taxa studied

Blainville's beaked whale.

Methods

Passive acoustic data were collected at 76 sites and 150 cumulative years of data were analysed to extract beaked whale echolocation clicks. Using an automated detector and subsequent weighted network clustering on spectral content and interclick interval of clicks, we determined the properties of a primary cluster of clicks with similar characteristics per site. These were compared within regions and across ocean basins and evaluated for suitability as population-level indicators.

Results

Spectral averages obtained from primary clusters of echolocation clicks identified at each site were similar in overall shape but varied in peak frequency by up to 8 kHz. We identified a latitudinal cline, with higher peak frequencies occurring in lower latitudes.

Main conclusions

It may be possible to acoustically delineate populations of Blainville's beaked whales. The documented negative correlation between signal peak frequency and latitude could relate to body size. Body size has been shown to influence signal frequency, with lower frequencies produced by larger animals, which are subsequently more common in higher latitudes for some species, although data are lacking to adequately investigate this for beaked whales. Prey size and depth may shape frequency content of echolocation signals, and larger prey items may occur in higher latitudes, resulting in lower signal frequencies of their predators.     

A review of big data analysis methods for baleen whale passive acoustic monitoring

Many organizations collect large passive acoustic monitoring (PAM) data sets that need to be efficiently and reliably analyzed. To determine appropriate methods for effective analysis of big PAM data sets, we undertook a literature review of baleen whale PAM analysis methods. Methodologies from 166 studies (published between 2000–2019) were summarized, and a detailed review was performed on the 94 studies that recorded more than 1,000 hr of acoustic data (“big data”). Analysis techniques for extracting baleen whale information from PAM data sets varied depending on the research observed. A spectrum of methodologies was used and ranged from manual analysis of all acoustic data by human experts to completely automated techniques with no manual validation. Based on this assessment, recommendations are provided to encourage robust research methods that are comparable across studies and sectors, achievable across research groups, and consistent with previous work. These include using automated techniques when possible to increase efficiency and repeatability, supplementing automation with manual review to calculate automated detector performance, and increasing consistency in terminology and presentation of results. This work can be used to facilitate discussion for minimum standards and best practices to be implemented in the field of marine mammal PAM.     

中国大陆海域沿岸搁浅和误捕的喙鲸类调查

从2009 年1 月至2010 年2 月,我们对中国大陆海域沿岸搁浅和误捕的喙鲸类头骨标本进行了测量和鉴
定。目前中国大陆海域搁浅和误捕的喙鲸类有5 种,分别为柯氏喙鲸、贝氏喙鲸、朗氏喙鲸、柏氏中喙鲸和小
中喙鲸,其中小中喙鲸为中国海域(包括台湾)的首次报道,也是在西北太平洋区的首次记录。小中喙鲸的主
要形态鉴别特征为有1 对纺锤形牙齿着生在下颌骨联合后面。经对已报道过的大陆海域搁浅和误捕的3 个雌性
银杏齿中喙鲸头骨标本和台湾省搁浅银杏齿中喙鲸的颅骨、鼻骨、下颌骨以及牙齿特征对比重新鉴定,订正大
陆的3 个样本为雌性柏氏中喙鲸。本文报道了大陆沿岸海域搁浅和误捕的喙鲸类最新调查研究结果。     

Passive acoustic density estimation of sperm whales in the Tongue of the Ocean,Bahamas

Long‐term passive acoustic monitoring of marine mammals on navy ranges provides the opportunity to better understand the potential impact of sonar on populations. The navy range in Tongue of the Ocean (TOTO), Bahamas contains extensive hydrophone arrays, potentially allowing estimation of the density of deep diving, vocally active species such as the sperm whale (Physeter macrocephalus). Previous visual surveys in TOTO have been of limited spatio–temporal coverage and resulted in only sporadic sightings of sperm whales, whereas passive acoustic observations suggest the species is present year round. However, until now the means of acoustically determining the specific number of individuals in each cluster has been limited. We used recently developed algorithms to identify the number of echolocating whales present during a 42 d study period. We screened a 297 h acoustic data set to determine the proportion of time animals were present; fifty 10 min samples during presence were analyzed to estimate the number of individuals vocalizing during each sample. These counts were combined with an independent estimate of the proportion of 10 min periods when tagged animals vocalize. The estimated average density was 0.16 whales/1,000 km² (CV 27%; 95% CI 0.095–0.264). The method is potentially applicable to other areas containing dense hydrophone arrays.     

Adult male Cuvier's beaked whales (Ziphius cavirostris) engage in prolonged bouts of synchronous diving

Studies of the social behavior of Cuvier's beaked whales (Ziphius cavirostris) are challenging because of their deep-water habitat usually far from shore and the limited time they spend at the surface. The sociality of these deepest diving mammals is of interest, however, especially for our understanding of how social systems evolve in extreme habitats. High levels of scarring suggest that males compete agonistically for access to females and so we predicted that associations among adult males would be unstable due to competitive exclusion. We tested this prediction by evaluating the diving behavior of animals within social groups off Cape Hatteras, North Carolina, considering diving synchrony a proxy for group membership. Using data from satellite-linked depth-recording tags, we found that adult male–male pairs showed extended periods of synchrony in diving behavior, while all pairs that included an adult male with an individual of another age and/or sex dove synchronously for less than a day. We assessed three hypotheses to explain these surprising results: sexual segregation; extended bouts of male–male competition; and the presence of male alliances. Finally, we considered testable predictions to distinguish among these explanations.     

Odontocete occurrence in relation to changes in oceanography at a remote equatorial Pacific seamount

Seamounts are considered hot spots of biodiversity and can aggregate pelagic predators and their prey. Passive acoustic monitoring was conducted over 3 mo in 2012 to document the occurrence of odontocetes near a seamount chain in the central equatorial Pacific in relation to oceanographic changes over time. Beaked whale echolocation signals were most frequently encountered. The main beaked whale signal was an unknown type, BW38, which resembled signals produced by Blainville's beaked whales. It had high occurrence during high sea surface temperature and low sea surface salinity. Cuvier's beaked whales were the second most detected. They had an opposite pattern and were encountered more often when sea surface temperature was low and net primary productivity was high. Risso's dolphins and short‐finned pilot whales had high acoustic densities, and echolocated predominantly at night. Risso's dolphins occurred more often during low sea surface height deviation. False killer whales were less frequently detected and mostly occurred during the day. Sperm whale detections were fewer than expected and associated with high chlorophyll a. Short duration Kogiidae encounters occurred on average every third day. These types of long‐term site studies are an informative tool to comparatively assess species composition, relative abundance, and relationship to oceanographic changes.     

Occurrence and social structure of Baird's beaked whales,Berardius bairdii,in the Commander Islands,Russia

The social structure of Baird's beaked whales is completely unstudied, and it is unknown if either females or males form long‐term social associations or occur in stable groups. In this paper we summarize our observations of individually identified animals over the span of 6 yr to provide insight on their long‐term social structure. We have identified 122 whales, with 28 of them encountered three times or more and thus included in the analysis of social structure. We found that the whales exhibited nonrandom patterns of social associations with some individuals preferentially associating with each other. Whales with more scarred skin had higher maximum association coefficients, which indicates that older animals and/or males were more inclined to form stable associations. Cluster analysis with a modularity test for gregariousness divided the whales into four clusters. Whales from the same clusters did not always occur together, but some individuals retained stable associations over several years. The strength of social relationships decayed over periods of months, with between‐year relationships showing little deviation from what would be expected if association was random. Generally these findings do not correspond to a stable society with fixed groups but instead suggest a fission‐fusion society with some stable alliances.     

Passive acoustic monitoring of the distribution patterns of Irrawaddy dolphins (Orcaella brevirostris) in the middle reaches of the Ayeyarwady River,Myanmar

The Irrawaddy dolphins (Orcaella brevirostris) are an endangered species. Thus, up-to-date information on the distribution pattern of dolphins is critical for its proper management and conservation. Using a towed passive acoustic monitoring device, the distribution pattern of the Irrawaddy dolphins in the middle reaches of the Ayeyarwady River, Myanmar, was investigated during a vessel-based survey between Mingun and Katha. This region was successively divided into segments 1–4 from upstream to downstream. Sixteen echolocation encounters, with a series of click trains separated by <8 min and 26 dolphin acoustic trajectories were recorded. The mean dolphin detection rate (animals/kilometer) across the four segments progressively increased from upstream to downstream. High relative abundance was observed in segment 4 (46%) and segment 1 (23%) which was consistent with findings from historical boat-based visual surveys. The averaged interclick intervals of each click train in segment 2 and 4 was significantly shorter than that in segment 3, indicating that the dolphins in these segments frequently use shorter-ranged biosonar. More frequent and consistent surveys with a systematic sampling track design that incorporates other factors and covering the whole distribution range along the Ayeyarwady River and at varied water levels are needed in the future.     

Estimating the number of whales entering trade using DNA profiling and capture-recapture analysis of market products

Surveys of commercial markets combined with molecular taxonomy (i.e. molecular monitoring) provide a means to detect products from illegal, unregulated and/or unreported (IUU) exploitation, including the sale of fisheries bycatch and wild meat (bushmeat). Capture-recapture analyses of market products using DNA profiling have the potential to estimate the total number of individuals entering the market. However, these analyses are not directly analogous to those of living individuals because a 'market individual' does not die suddenly but, instead, remains available for a time in decreasing quantities, rather like the exponential decay of a radioactive isotope. Here we use mitochondrial DNA (mtDNA) sequences and microsatellite genotypes to individually identify products from North Pacific minke whales (Balaenoptera acutorostrata ssp.) purchased in 12 surveys of markets in the Republic of (South) Korea from 1999 to 2003. By applying a novel capture-recapture model with a decay rate parameter to the 205 unique DNA profiles found among 289 products, we estimated that the total number of whales entering trade across the five-year survey period was 827 (SE, 164; CV, 0.20) and that the average 'half-life' of products from an individual whale on the market was 1.82 months (SE, 0.24; CV, 0.13). Our estimate of whales in trade (reflecting the true numbers killed) was significantly greater than the officially reported bycatch of 458 whales for this period. This unregulated exploitation has serious implications for the survival of this genetically distinct coastal population. Although our capture-recapture model was developed for specific application to the Korean whale-meat markets, the exponential decay function could be modified to improve the estimates of trade in other wildmeat or fisheries markets or abundance of living populations by noninvasive genotyping.     

Humpback whale singing activity off the Goan coast in the Eastern Arabian Sea

For over two decades, passive acoustic monitoring (PAM) methods have been successfully employed around the world for studying aquatic megafauna. PAM-driven studies in Indian waters have so far been relatively very scarce. Furthermore, cetacean populations inhabiting the north western Indian Ocean are far less studied than those in many other regions around the world. This work likely constitutes the first systematic study of the vocal repertoire of humpback whales (Megaptera novaeangliae) at a near-shore site along the western coast of India. Analysis of the observed vocalizations provides an insight into the behaviour of the species. This is significant as it assists in developing a better understanding of the habitat use of the non-migratory Arabian Sea humpback whale population. In contrast, other breeding populations such as those around the North Atlantic, South Pacific and Australia have been relatively well studied. Underwater passive acoustic data were collected during March 2017 using an autonomous logger at a shallow-water site off the eastern edge of Grande Island off the coast of Goa. Humpback whale vocalizations were found to occur over multiple days in the recordings. Time–frequency contours of individual units of vocalization were extracted with the aid of an automatic detection technique and the characteristics of the units were measured. Further, successive units were analysed for formation of phrases and themes. Reconstruction of putative songs from the identified units and themes was not possible due to the limitations imposed by the nature of data collection. Detailed analyses of units, phrases and themes are presented.     

No shallow talk: Cryptic strategy in the vocal communication of Blainville's beaked whales

Communicating animals must balance fitness benefits against the costs of signaling, such as increased predation risk. Cetaceans communicate mainly with sound and near‐surface vocalizations can place signalers at risk from shallow‐diving top‐predators with acute hearing such as killer whales. Beaked whales are deep divers living in small cohesive groups with little social defense from predation. Little if anything is known about their acoustic communication. Here, eight Blainville's beaked whales were studied with suction cup attached DTags to provide the first report on social communication as a function of diving behavior for any of the 21 ziphiid species. Tagged whales produced two previously unrecorded signals with apparent communicative functions: (1) fast series of ultrasonic frequency modulated clicks (rasps) were recorded from six individuals, and (2) harmonically rich short whistles with a mean fundamental frequency of 12 kHz were recorded from one whale at up to 900 m depth, the deepest whistles recorded from a marine mammal. Blainville's were silent 80% of the time, whenever shallower than 170 m depth and during the prolonged (19 min) silent ascents from vocal dives. This behavior limits the ability of shallow‐diving predators to track Blainville's acoustically and may provide a striking example of the evolutionary influence of the risk of predation on animal communication.     

Vocal activity of tropical dolphins is inhibited by the presence of killer whales,Orcinus orca

Research has suggested killer whale (Orcinus orca) predation may affect cetacean vocal behavior; however, few data exist to test this hypothesis. Data collected during 40,976 km of visual and acoustic shipboard surveys in the tropical Pacific Ocean, including 1,232 detections of 13 species, were examined to determine if changes in dolphin vocal activity could be attributed to the presence of killer whales. Generalized linear models and Random Forest analyses were used to test the hypothesis that dolphin vocal activity was related to the distance and time to the nearest killer whale sighting. Both results show that dolphin vocalizations were inversely correlated with the temporal proximity of killer whales (P < 0.05). Despite the relative rarity of killer whales in the tropics, they appear to influence vocal behavior of nearby dolphin schools. This disruption in communication may not significantly impact interactions necessary for survival in tropical waters where killer whale density is low. However, in temperate climates, where increased productivity supports a greater abundance of killer whales, this interruption in communication may have a greater impact. The lower incidence of whistling dolphins in temperate waters may be related to the greater abundance of killer whales in these areas.     

Handling dolphin detections from C-PODs,with the development of acoustic parameters for verification and the exploration of species identification possibilities

C-PODs are static passive acoustic monitoring devices used to detect odontocete vocalizations in the range of 20–160 kHz. However, falsely classified detections may be an issue, particularly with broadband species (i.e. many dolphin species) due to anthropogenic and other noise occurring at the same frequency. While porpoise detections are verified using species-specific acoustic parameters, the equivalent does not currently exist for verifying dolphin detections. Development of such parameters would increase the accuracy of dolphin detections and eliminate the need for additional monitoring techniques or devices, reducing the cost of monitoring programmes. Herein, we present parameters based on acoustic characteristics of bottlenose (n = 29), common (n = 19) and Risso’s (n = 99) dolphin click trains, sighted within 1 km of C-PODs during land-based surveys, for in-software verification. Overlap of click train parameters among dolphin species prevented robust species identification; therefore, parameters were devised for these dolphin species collectively using frequency, inter-click interval and click train duration. A data set of 4898 Detection Positive Hours was visually verified using these parameters. The temporal and spatial patterns in the visually verified data were similar to land-based observations, suggesting the parameters operate at an acceptable accuracy. However, 68% of high-, moderate- and low-quality KERNO detections were false-positive. Our results suggest that the accuracy of classifiers and quality class weightings are site-specific, and we highlight the importance of data exploration to make the most appropriate software choices based on the aims of a study.     

Review of the distribution of Mesoplodon species (order Cetacea, family Ziphiidae) in the North Atlantic

Four species of beaked whales of the genus Mesoplodon regularly occur in the North Atlantic. These are Sowerby's Beaked Whale Mesoplodon bidens, Blainville's Beaked Whale Mesoplodon densirostris, Gervais Beaked Whale Mesoplodon europaeus and True's Beaked Whale Mesoplodon mirus. These four species differ in their distribution within this area. M. bidens is the most northerly recorded species followed by M. mirus. M. densirostris and M. europaeus both generally occur further south and their distributions cross the equator. The distribution of Mesoplodon species may relate to variations in water temperature.