Click detection rate variability of central North Pacific sperm whales from passive acoustic towed arrays.

Passive acoustic monitoring (PAM) is an optimal method for detecting and monitoring cetaceans as they frequently produce sound while underwater. Cue counting, counting acoustic cues of deep-diving cetaceans instead of animals, is an alternative method for density estimation, but requires an average cue production rate to convert cue density to animal density. Limited information about click rates exists for sperm whales in the central North Pacific Ocean. In the absence of acoustic tag data, we used towed hydrophone array data to calculate the first sperm whale click rates from this region and examined their variability based on click type, location, distance of whales from the array, and group size estimated by visual observers. Our findings show click type to be the most important variable, with groups that include codas yielding the highest click rates. We also found a positive relationship between group size and click detection rates that may be useful for acoustic predictions of group size in future studies. Echolocation clicks detected using PAM methods are often the only indicator of deep-diving cetacean presence. Understanding the factors affecting their click rates provides important information for acoustic density estimation.

An unknown nocturnal call type in the Mariana Archipelago.

In spring/summer of 2018 and 2021, the Pacific Islands Fisheries Science Center Cetacean Research Program deployed drifting acoustic recorders in the U.S. Exclusive Economic Zones surrounding the Mariana Archipelago. Manual assessments revealed a low-frequency (median 473-554 Hz), short-duration (median 0.596 s), stereotypic tonal nocturnal call throughout the Mariana Archipelago. Based on time of year, spatiotemporal patterns, clear division among calls (i.e., no chorusing), comparisons with known vocalizations of whales, turtles, and fish, and presence of Bryde's whale calls, and because the call has not been detected elsewhere, we hypothesize this 500-Hz pulsed call is produced by Bryde's whales (Balaenoptera brydei).

Odontocete spatial patterns and temporal drivers of detection at sites in the Hawaiian islands.

Successful conservation and management of marine top predators rely on detailed documentation of spatiotemporal behavior. For cetacean species, this information is key to defining stocks, habitat use, and mitigating harmful interactions. Research focused on this goal is employing methodologies such as visual observations, tag data, and passive acoustic monitoring (PAM) data. However, many studies are temporally limited or focus on only one or few species. In this study, we make use of an existing long-term (2009-2019), labeled PAM data set to examine spatiotemporal patterning of at least 10 odontocete (toothed whale) species in the Hawaiian Islands using compositional analyses and modeling techniques. Species composition differs among considered sites, and this difference is robust to seasonal movement patterns. Temporally, hour of day was the most significant predictor of detection across species and sites, followed by season, though patterns differed among species. We describe long-term trends in species detection at one site and note that they are markedly similar for many species. These trends may be related to long-term, underlying oceanographic cycles that will be the focus of future study. We demonstrate the variability of temporal patterns even at relatively close sites, which may imply that wide-ranging models of species presence are missing key fine-scale movement patterns. Documented seasonal differences in detection also highlights the importance of considering season in survey design both regionally and elsewhere. We emphasize the utility of long-term, continuous monitoring in highlighting temporal patterns that may relate to underlying climatic states and help us predict responses to climate change. We conclude that long-term PAM records are a valuable resource for documenting spatiotemporal patterns and can contribute many insights into the lives of top predators, even in highly studied regions such as the Hawaiian Islands.

Evidence from sperm whale clans of symbolic marking in non-human cultures.

Culture, a pillar of the remarkable ecological success of humans, is increasingly recognized as a powerful force structuring nonhuman animal populations. A key gap between these two types of culture is quantitative evidence of symbolic markers-seemingly arbitrary traits that function as reliable indicators of cultural group membership to conspecifics. Using acoustic data collected from 23 Pacific Ocean locations, we provide quantitative evidence that certain sperm whale acoustic signals exhibit spatial patterns consistent with a symbolic marker function. Culture segments sperm whale populations into behaviorally distinct clans, which are defined based on dialects of stereotyped click patterns (codas). We classified 23,429 codas into types using contaminated mixture models and hierarchically clustered coda repertoires into seven clans based on similarities in coda usage; then we evaluated whether coda usage varied with geographic distance within clans or with spatial overlap between clans. Similarities in within-clan usage of both "identity codas" (coda types diagnostic of clan identity) and "nonidentity codas" (coda types used by multiple clans) decrease as space between repertoire recording locations increases. However, between-clan similarity in identity, but not nonidentity, coda usage decreases as clan spatial overlap increases. This matches expectations if sympatry is related to a measurable pressure to diversify to make cultural divisions sharper, thereby providing evidence that identity codas function as symbolic markers of clan identity. Our study provides quantitative evidence of arbitrary traits, resembling human ethnic markers, conveying cultural identity outside of humans, and highlights remarkable similarities in the distributions of human ethnolinguistic groups and sperm whale clans.

Discriminating and classifying odontocete echolocation clicks in the Hawaiian Islands using machine learning methods.

Passive acoustic monitoring (PAM) has proven a powerful tool for the study of marine mammals, allowing for documentation of biologically relevant factors such as movement patterns or animal behaviors while remaining largely non-invasive and cost effective. From 2008-2019, a set of PAM recordings covering the frequency band of most toothed whale (odontocete) echolocation clicks were collected at sites off the islands of Hawai'i, Kaua'i, and Pearl and Hermes Reef. However, due to the size of this dataset and the complexity of species-level acoustic classification, multi-year, multi-species analyses had not yet been completed. This study shows how a machine learning toolkit can effectively mitigate this problem by detecting and classifying echolocation clicks using a combination of unsupervised clustering methods and human-mediated analyses. Using these methods, it was possible to distill ten unique echolocation click 'types' attributable to regional odontocetes at the genus or species level. In one case, auxiliary sightings and recordings were used to attribute a new click type to the rough-toothed dolphin, Steno bredanensis. Types defined by clustering were then used as input classes in a neural-network based classifier, which was trained, tested, and evaluated on 5-minute binned data segments. Network precision was variable, with lower precision occurring most notably for false killer whales, Pseudorca crassidens, across all sites (35-76%). However, accuracy and recall were high (>96% and >75%, respectively) in all cases except for one type of short-finned pilot whale, Globicephala macrorhynchus, call class at Kaua'i and Pearl and Hermes Reef (recall >66%). These results emphasize the utility of machine learning in analysis of large PAM datasets. The classifier and timeseries developed here will facilitate further analyses of spatiotemporal patterns of included toothed whales. Broader application of these methods may improve the efficiency of global multi-species PAM data processing for echolocation clicks, which is needed as these datasets continue to grow.

Model-based localization of deep-diving cetaceans using towed line array acoustic data.

Passive acoustic monitoring using a towed line array of hydrophones is a standard method for localizing cetaceans during line-transect cetacean abundance surveys. Perpendicular distances estimated between localized whales and the trackline are essential for abundance estimation using acoustic data. Uncertainties in the acoustic data from hydrophone movement, sound propagation effects, errors in the time of arrival differences, and whale depth are not accounted for by most two-dimensional localization methods. Consequently, location and distance estimates for deep-diving cetaceans may be biased, creating uncertainty in abundance estimates. Here, a model-based localization approach is applied to towed line array acoustic data that incorporates sound propagation effects, accounts for sources of error, and localizes in three dimensions. The whale's true distance, ship trajectory, and whale movement greatly affected localization results in simulations. The localization method was applied to real acoustic data from two separate sperm whales, resulting in three-dimensional distance and depth estimates with position bounds for each whale. By incorporating sources of error, this three-dimensional model-based approach provides a method to address and integrate the inherent uncertainties in towed array acoustic data for more robust localization.

Acoustic classification of false killer whales in the Hawaiian islands based on comprehensive vocal repertoire.

Use of underwater passive acoustic datasets for species-specific inference requires robust classification systems to identify encounters to species from characteristics of detected sounds. A suite of routines designed to efficiently detect cetacean sounds, extract features, and classify the detection to species is described using ship-based, visually verified detections of false killer whales (Pseudorca crassidens). The best-performing model included features from clicks, whistles, and burst pulses, which correctly classified 99.6% of events. This case study illustrates use of these tools to build classifiers for any group of cetacean species and assess classification confidence when visual confirmation is not available.

Co-occurrence of beaked whale strandings and naval sonar in the Mariana Islands, Western Pacific.

Mid-frequency active sonar (MFAS), used for antisubmarine warfare (ASW), has been associated with multiple beaked whale (BW) mass stranding events. Multinational naval ASW exercises have used MFAS offshore of the Mariana Archipelago semi-annually since 2006. We report BW and MFAS acoustic activity near the islands of Saipan and Tinian from March 2010 to November 2014. Signals from Cuvier's (Ziphius cavirostris) and Blainville's beaked whales (Mesoplodon densirostris), and a third unidentified BW species, were detected throughout the recording period. Both recorders documented MFAS on 21 August 2011 before two Cuvier's beaked whales stranded on 22-23 August 2011. We compared the history of known naval operations and BW strandings from the Mariana Archipelago to consider potential threats to BW populations. Eight BW stranding events between June 2006 and January 2019 each included one to three animals. Half of these strandings occurred during or within 6 days after naval activities, and this co-occurrence is highly significant. We highlight strandings of individual BWs can be associated with ASW, and emphasize the value of ongoing passive acoustic monitoring, especially for beaked whales that are difficult to visually detect at sea. We strongly recommend more visual monitoring efforts, at sea and along coastlines, for stranded cetaceans before, during and after naval exercises.

Oceanographic barriers, divergence, and admixture: Phylogeography and taxonomy of two putative subspecies of short-finned pilot whale.

Genomic phylogeography plays an important role in describing evolutionary processes and their geographic, ecological, or cultural drivers. These drivers are often poorly understood in marine environments, which have fewer obvious barriers to mixing than terrestrial environments. Taxonomic uncertainty of some taxa (e.g., cetaceans), due to the difficulty in obtaining morphological data, can hamper our understanding of these processes. One such taxon, the short-finned pilot whale, is recognized as a single global species but includes at least two distinct morphological forms described from stranding and drive hunting in Japan, the "Naisa" and "Shiho" forms. Using samples (n = 735) collected throughout their global range, we examine phylogeographic patterns of divergence by comparing mitogenomes and nuclear SNP loci. Our results suggest three types within the species: an Atlantic Ocean type, a western/central Pacific and Indian Ocean (Naisa) type, and an eastern Pacific Ocean and northern Japan (Shiho) type. mtDNA control region differentiation indicates these three types form two subspecies, separated by the East Pacific Barrier: Shiho short-finned pilot whale, in the eastern Pacific Ocean and northern Japan, and Naisa short-finned pilot whale, throughout the remainder of the species' distribution. Our data further indicate two diverging populations within the Naisa subspecies, in the Atlantic Ocean and western/central Pacific and Indian Oceans, separated by the Benguela Barrier off South Africa. This study reveals a process of divergence and speciation within a globally-distributed, mobile marine predator, and indicates the importance of the East Pacific Barrier to this evolutionary process.

Context-dependent variability in blue whale acoustic behaviour.

Acoustic communication is an important aspect of reproductive, foraging and social behaviours for many marine species. Northeast Pacific blue whales (Balaenoptera musculus) produce three different call types-A, B and D calls. All may be produced as singular calls, but A and B calls also occur in phrases to form songs. To evaluate the behavioural context of singular call and phrase production in blue whales, the acoustic and dive profile data from tags deployed on individuals off southern California were assessed using generalized estimating equations. Only 22% of all deployments contained sounds attributed to the tagged animal. A larger proportion of tagged animals were female (47%) than male (13%), with 40% of unknown sex. Fifty per cent of tags deployed on males contained sounds attributed to the tagged whale, while only a few (5%) deployed on females did. Most calls were produced at shallow depths (less than 30 m). Repetitive phrasing (singing) and production of singular calls were most common during shallow, non-lunging dives, with the latter also common during surface behaviour. Higher sound production rates occurred during autumn than summer and they varied with time-of-day: singular call rates were higher at dawn and dusk, while phrase production rates were highest at dusk and night.

Familial social structure and socially driven genetic differentiation in Hawaiian short-finned pilot whales.

Social structure can have a significant impact on divergence and evolution within species, especially in the marine environment, which has few environmental boundaries to dispersal. On the other hand, genetic structure can affect social structure in many species, through an individual preference towards associating with relatives. One social species, the short-finned pilot whale (Globicephala macrorhynchus), has been shown to live in stable social groups for periods of at least a decade. Using mitochondrial control sequences from 242 individuals and single nucleotide polymorphisms from 106 individuals, we examine population structure among geographic and social groups of short-finned pilot whales in the Hawaiian Islands, and test for links between social and genetic structure. Our results show that there are at least two geographic populations in the Hawaiian Islands: a Main Hawaiian Islands (MHI) population and a Northwestern Hawaiian Islands/Pelagic population (FST and ΦST p < .001), as well as an eastern MHI community and a western MHI community (FST p = .009). We find genetically driven social structure, or high relatedness among social units and clusters (p < .001), and a positive relationship between relatedness and association between individuals (p < .0001). Further, socially organized clusters are genetically distinct, indicating that social structure drives genetic divergence within the population, likely through restricted mate selection (FST p = .05). This genetic divergence among social groups can make the species less resilient to anthropogenic or ecological disturbance. Conservation of this species therefore depends on understanding links among social structure, genetic structure and ecological variability within the species.

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.

Fin whale song variability in southern California and the Gulf of California.

Songs are distinct, patterned sounds produced by a variety of animals including baleen whales. Fin whale songs, which consist of short pulses repeated at regular interpulse intervals (IPIs), have been suggested as a tool to distinguish populations. Fin whale songs were analyzed from data collected from 2000-2012 in Southern California and from 2004-2010 in the Gulf of California using autonomous acoustic recorders. IPIs were measured for each identifiable song sequence during two random days of each month with recordings. Four distinct song types were identified: long doublet, short doublet, long triplet, and short triplet. Long and short doublets were the dominant songs in Southern California, while long and short triplets were dominant in the Gulf of California. An abrupt change in song type occurred in both areas during the monitoring period. We argue that each song type is unique to a population and these changes represent a shift in the primary population in the monitoring area. Occasional temporal and spatial song overlap indicated some exchange or visitation among populations. Fin whales appear to synchronize and gradually modify song rhythm over long time scales. A better understanding of the evolutionary and ecological importance of songs to fin whale populations is needed.

Limited trophic partitioning among sympatric delphinids off a tropical oceanic atoll.

Understanding trophic relationships among marine predators in remote environments is challenging, but it is critical to understand community structure and dynamics. In this study, we used stable isotope analysis of skin biopsies to compare the isotopic, and thus, trophic niches of three sympatric delphinids in the waters surrounding Palmyra Atoll, in the Central Tropical Pacific: the melon-headed whale (Peponocephala electra), Gray's spinner dolphin (Stenella longirostris longirostris), and the common bottlenose dolphin (Tursiops truncatus). δ15N values suggested that T. truncatus occupied a significantly higher trophic position than the other two species. δ13C values did not significantly differ between the three delphinds, potentially indicating no spatial partitioning in depth or distance from shore in foraging among species. The dietary niche area-determined by isotopic variance among individuals-of T. truncatus was also over 30% smaller than those of the other species taken at the same place, indicating higher population specialization or lower interindividual variation. For P. electra only, there was some support for intraspecific variation in foraging ecology across years, highlighting the need for temporal information in studying dietary niche. Cumulatively, isotopic evidence revealed surprisingly little evidence for trophic niche partitioning in the delphinid community of Palmyra Atoll compared to other studies. However, resource partitioning may happen via other behavioral mechanisms, or prey abundance or availability may be adequate to allow these three species to coexist without any such partitioning. It is also possible that isotopic signatures are inadequate to detect trophic partitioning in this environment, possibly because isotopes of prey are highly variable or insufficiently resolved to allow for differentiation.

Discrimination of frequency-modulated Baleen whale downsweep calls with overlapping frequencies.

Automatic classification of fin, sei, and blue whale frequency modulated downsweeps has been a challenging task for bioacousticians. These calls overlap in frequency range and have similar time durations. The traditional spectrogram methodology, the Short Time Fourier Transform, tends to be ineffective because of the large temporal ambiguities needed to achieve the necessary frequency resolution to study the fine time-frequency (TF) structures. Spectrograms generated with the Pseudo Wigner-Ville Distribution (PWVD) provide much higher simultaneous TF resolution compared with the traditional method. The PWVD allows bioacousticians to study the fine TF structures of the sound, such as the instantaneous frequency, instantaneous bandwidth, contour slope, etc. These features set the foundation of identifying sounds that are usually considered difficult to discriminate using the traditional method. Wigner-Ville distribution of the baleen whale downsweeps showed distinguishable characteristics; for example, the TF contour of fin and sei whales exhibited concave and convex shapes, which have never been reported in the literature. A Support Vector Machine classifier was trained and tested based on the parameters extracted from the PWVD.

Synchronous seasonal change in fin whale song in the North Pacific.

Fin whale (Balaenoptera physalus) song consists of down-swept pulses arranged into stereotypic sequences that can be characterized according to the interval between successive pulses. As in blue (B. musculus) and humpback whales (Megaptera novaeangliae), these song sequences may be geographically distinct and may correlate with population boundaries in some regions. We measured inter-pulse intervals of fin whale songs within year-round acoustic datasets collected between 2000 and 2006 in three regions of the eastern North Pacific: Southern California, the Bering Sea, and Hawaii. A distinctive song type that was recorded in all three regions is characterized by singlet and doublet inter-pulse intervals that increase seasonally, then annually reset to the same shorter intervals at the beginning of each season. This song type was recorded in the Bering Sea and off Southern California from September through May and off Hawaii from December through April, with the song interval generally synchronized across all monitoring locations. The broad geographic and seasonal occurrence of this particular fin whale song type may represent a single population broadly distributed throughout the eastern Pacific with no clear seasonal migratory pattern. Previous studies attempting to infer population structure of fin whales in the North Pacific using synchronous individual song samples have been unsuccessful, likely because they did not account for the seasonal lengthening in song intervals observed here.

Estimating historical eastern North Pacific blue whale catches using spatial calling patterns.

Blue whales (Balaenoptera musculus) were exploited extensively around the world and remain endangered. In the North Pacific their population structure is unclear and current status unknown, with the exception of a well-studied eastern North Pacific (ENP) population. Despite existing abundance estimates for the ENP population, it is difficult to estimate pre-exploitation abundance levels and gauge their recovery because historical catches of the ENP population are difficult to separate from catches of other populations in the North Pacific. We collated previously unreported Soviet catches and combined these with known catches to form the most current estimates of North Pacific blue whale catches. We split these conflated catches using recorded acoustic calls from throughout the North Pacific, the knowledge that the ENP population produces a different call than blue whales in the western North Pacific (WNP). The catches were split by estimating spatiotemporal occurrence of blue whales with generalized additive models fitted to acoustic call patterns, which predict the probability a catch belonged to the ENP population based on the proportion of calls of each population recorded by latitude, longitude, and month. When applied to the conflated historical catches, which totaled 9,773, we estimate that ENP blue whale catches totaled 3,411 (95% range 2,593 to 4,114) from 1905-1971, and amounted to 35% (95% range 27% to 42%) of all catches in the North Pacific. Thus most catches in the North Pacific were for WNP blue whales, totaling 6,362 (95% range 5,659 to 7,180). The uncertainty in the acoustic data influence the results substantially more than uncertainty in catch locations and dates, but the results are fairly insensitive to the ecological assumptions made in the analysis. The results of this study provide information for future studies investigating the recovery of these populations and the impact of continuing and future sources of anthropogenic mortality.

Nuclear and mitochondrial patterns of population structure in North Pacific false killer whales (Pseudorca crassidens).

False killer whales (Pseudorca crassidens) are large delphinids typically found in deep water far offshore. However, in the Hawaiian Archipelago, there are 2 resident island-associated populations of false killer whales, one in the waters around the main Hawaiian Islands (MHI) and one in the waters around the Northwestern Hawaiian Islands (NWHI). We use mitochondrial DNA (mtDNA) control region sequences and genotypes from 16 nuclear DNA (nucDNA) microsatellite loci from 206 individuals to examine levels of differentiation among the 2 island-associated populations and offshore animals from the central and eastern North Pacific. Both mtDNA and nucDNA exhibit highly significant differentiation between populations, confirming limited gene flow in both sexes. The mtDNA haplotypes exhibit a strong pattern of phylogeographic concordance, with island-associated populations sharing 3 closely related haplotypes not found elsewhere in the Pacific. However, nucDNA data suggest that NWHI animals are at least as differentiated from MHI animals as they are from offshore animals. The patterns of differentiation revealed by the 2 marker types suggest that the island-associated false killer whale populations likely share a common colonization history, but have limited contemporary gene flow.

Accounting for subgroup structure in line-transect abundance estimates of false killer whales (Pseudorca crassidens) in Hawaiian waters.

For biological populations that form aggregations (or clusters) of individuals, cluster size is an important parameter in line-transect abundance estimation and should be accurately measured. Cluster size in cetaceans has traditionally been represented as the total number of individuals in a group, but group size may be underestimated if group members are spatially diffuse. Groups of false killer whales (Pseudorca crassidens) can comprise numerous subgroups that are dispersed over tens of kilometers, leading to a spatial mismatch between a detected group and the theoretical framework of line-transect analysis. Three stocks of false killer whales are found within the U.S. Exclusive Economic Zone of the Hawaiian Islands (Hawaiian EEZ): an insular main Hawaiian Islands stock, a pelagic stock, and a Northwestern Hawaiian Islands (NWHI) stock. A ship-based line-transect survey of the Hawaiian EEZ was conducted in the summer and fall of 2010, resulting in six systematic-effort visual sightings of pelagic (n = 5) and NWHI (n = 1) false killer whale groups. The maximum number and spatial extent of subgroups per sighting was 18 subgroups and 35 km, respectively. These sightings were combined with data from similar previous surveys and analyzed within the conventional line-transect estimation framework. The detection function, mean cluster size, and encounter rate were estimated separately to appropriately incorporate data collected using different methods. Unlike previous line-transect analyses of cetaceans, subgroups were treated as the analytical cluster instead of groups because subgroups better conform to the specifications of line-transect theory. Bootstrap values (n = 5,000) of the line-transect parameters were randomly combined to estimate the variance of stock-specific abundance estimates. Hawai'i pelagic and NWHI false killer whales were estimated to number 1,552 (CV = 0.66; 95% CI = 479-5,030) and 552 (CV = 1.09; 95% CI = 97-3,123) individuals, respectively. Subgroup structure is an important factor to consider in line-transect analyses of false killer whales and other species with complex grouping patterns.

Spatio-temporal patterns of beaked whale echolocation signals in the North Pacific.

At least ten species of beaked whales inhabit the North Pacific, but little is known about their abundance, ecology, and behavior, as they are elusive and difficult to distinguish visually at sea. Six of these species produce known species-specific frequency modulated (FM) echolocation pulses: Baird's, Blainville's, Cuvier's, Deraniyagala's, Longman's, and Stejneger's beaked whales. Additionally, one described FM pulse (BWC) from Cross Seamount, Hawai'i, and three unknown FM pulse types (BW40, BW43, BW70) have been identified from almost 11 cumulative years of autonomous recordings at 24 sites throughout the North Pacific. Most sites had a dominant FM pulse type with other types being either absent or limited. There was not a strong seasonal influence on the occurrence of these signals at any site, but longer time series may reveal smaller, consistent fluctuations. Only the species producing BWC signals, detected throughout the Pacific Islands region, consistently showed a diel cycle with nocturnal foraging. By comparing stranding and sighting information with acoustic findings, we hypothesize that BWC signals are produced by ginkgo-toothed beaked whales. BW43 signal encounters were restricted to Southern California and may be produced by Perrin's beaked whale, known only from Californian waters. BW70 signals were detected in the southern Gulf of California, which is prime habitat for Pygmy beaked whales. Hubb's beaked whale may have produced the BW40 signals encountered off central and southern California; however, these signals were also recorded off Pearl and Hermes Reef and Wake Atoll, which are well south of their known range.