Annual phenology and migration routes to breeding grounds in western-central North Pacific sei whales.

The sei whale (Balaenoptera borealis) is an important species among baleen whales in the North Pacific and plays a significant role in the ecosystem. Despite the importance of this species, information regarding its migration patterns and breeding locations remains limited. To enhance the understanding of the phenology of North Pacific sei whales, we deployed satellite-monitored tags on these whales in the western and central North Pacific from 2017 to 2023. We fitted 55 sei whale tracks to a state-space model to describe the whales' seasonal movements at feeding grounds and their migratory behavior. The whales typically leave their feeding grounds between November and December, with migration pathways extending from off Japan to the west of the Hawaiian Islands. These southward transits converge in the waters of the Marshall Islands and north of Micronesia between 20° N and 7° N, which appear to be breeding grounds. After a brief stay at these breeding grounds, the whales migrate northward from January to February, reaching their feeding grounds around 30°N by March. To the best of our knowledge, this is the first study to present the phenology of feeding and breeding seasons and the migration pattern of North Pacific sei whales.

Localization of peptide hormones in the placentas of Bryde's (Balaenoptera brydei), sei (B. borealis), and common minke (B. acutorostrata) whales.

In this study, we examined the morphological features of the placentas from 3 species of rorqual whales (Balaenopteridae), namely Bryde's (Balaenoptera brydei), sei (B. borealis), and common minke (B. acutorostrata) whales, and verified the secretion of 2 placental-specific peptide hormones, placental lactogen (PL) and chorionic gonadotropin (CG). The placentas were collected in the second phase of the Japanese Whale Research Program under a special permit in the North Pacific (JARPN II) between 2009 and 2010. For all three species of rorqual whales, as the fetus grew, the interdigitation between the maternal endometrial folds and chorionic villi became more complicated, and many blood capillaries of chorionic villi and endometrium became larger and infiltrated the trophoblast cells and endometrial epithelial cells, respectively. In the immunohistochemical examination, the trophoblast cells (except for areolar trophoblast cells) showed immunoreactivities for the PL and luteinizing hormone (LH) antibodies, and this phenomenon was similar in the placentas of all 3 rorqual whale species. Our results suggest that PL and LH-like CG play roles in regulating pregnancy in the placenta of cetacean.

An accelerometer-derived ballistocardiogram method for detecting heart rate in free-ranging marine mammals.

Physio-logging methods, which use animal-borne devices to record physiological variables, are entering a new era driven by advances in sensor development. However, existing datasets collected with traditional bio-loggers, such as accelerometers, still contain untapped eco-physiological information. Here, we present a computational method for extracting heart rate from high-resolution accelerometer data using a ballistocardiogram. We validated our method with simultaneous accelerometer-electrocardiogram tag deployments in a controlled setting on a killer whale (Orcinus orca) and demonstrate the predictions correspond with previously observed cardiovascular patterns in a blue whale (Balaenoptera musculus), including the magnitude of apneic bradycardia and increase in heart rate prior to and during ascent. Our ballistocardiogram method may be applied to mine heart rates from previously collected accelerometery data and expand our understanding of comparative cardiovascular physiology.

Update on frequency decline of Northeast Pacific blue whale (Balaenoptera musculus) calls.

Worldwide, the frequency (pitch) of blue whale (Balaenoptera musculus) calls has been decreasing since first recorded in the 1960s. This frequency decline occurs over annual and inter-annual timescales and has recently been documented in other baleen whale species, yet it remains unexplained. In the Northeast Pacific, blue whales produce two calls, or units, that, when regularly repeated, are referred to as song: A and B calls. In this population, frequency decline has thus far only been examined in B calls. In this work, passive acoustic data collected in the Southern California Bight from 2006 to 2019 were examined to determine if A calls are also declining in frequency and whether the call pulse rate was similarly impacted. Additionally, frequency measurements were made for B calls to determine whether the rate of frequency decline is the same as was calculated when this phenomenon was first reported in 2009. We found that A calls decreased at a rate of 0.32 Hz yr-1 during this period and that B calls were still decreasing, albeit at a slower rate (0.27 Hz yr-1) than reported previously. The A call pulse rate also declined over the course of the study, at a rate of 0.006 pulses/s yr-1. With this updated information, we consider the various theories that have been proposed to explain frequency decline in blue whales. We conclude that no current theory adequately accounts for all aspects of this phenomenon and consider the role that individual perception of song frequency may play. To understand the cause behind call frequency decline, future studies might want to explore the function of these songs and the mechanism for their synchronization. The ubiquitous nature of the frequency shift phenomenon may indicate a consistent level of vocal plasticity and fine auditory processing abilities across baleen whale species.

A method for tracking blue whales (Balaenoptera musculus) with a widely spaced network of ocean bottom seismometers.

Passive acoustic monitoring is an important tool for studying marine mammals. Ocean bottom seismometer networks provide data sets of opportunity for studying blue whales (Balaenoptera musculus) which vocalize extensively at seismic frequencies. We describe methods to localize calls and obtain tracks using the B call of northeast Pacific blue whale recorded by a large network of widely spaced ocean bottom seismometers off the coast of the Pacific Northwest. The first harmonic of the B call at ~15 Hz is detected using spectrogram cross-correlation. The seasonality of calls, inferred from a dataset of calls identified by an analyst, is used to estimate the probability that detections are true positives as a function of the strength of the detection. Because the spacing of seismometers reaches 70 km, faint detections with a significant probability of being false positives must be considered in multi-station localizations. Calls are located by maximizing a likelihood function which considers each strong detection in turn as the earliest arrival time and seeks to fit the times of detections that follow within a feasible time and distance window. An alternative procedure seeks solutions based on the detections that maximize their sum after weighting by detection strength and proximity. Both approaches lead to many spurious solutions that can mix detections from different B calls and include false detections including misidentified A calls. Tracks that are reliable can be obtained iteratively by assigning detections to localizations that are grouped in space and time, and requiring groups of at least 20 locations. Smooth paths are fit to tracks by including constraints that minimize changes in speed and direction while fitting the locations to their uncertainties or applying the double difference relocation method. The reliability of localizations for future experiments might be improved by increasing sampling rates and detecting harmonics of the B call.

Modeling changes in baleen whale seasonal abundance, timing of migration, and environmental variables to explain the sudden rise in entanglements in California.

We document changes in the number of sightings and timing of humpback (Megaptera novaeangliae), blue (Balaenoptera musculus), and gray (Eschrichtius robustus) whale migratory phases in the vicinity of the Farallon Islands, California. We hypothesized that changes in the timing of migration off central California were driven by local oceanography, regional upwelling, and basin-scale climate conditions. Using 24 years of daily whale counts collected from Southeast Farallon Island, we developed negative binomial regression models to evaluate trends in local whale sightings over time. We then used linear models to assess trends in the timing of migration, and to identify potential environmental drivers. These drivers included local, regional and basin-scale patterns; the latter included the El Niño Southern Oscillation, the Pacific Decadal Oscillation, and the North Pacific Gyre Oscillation, which influence, wind-driven upwelling, and overall productivity in the California Current System. We then created a forecast model to predict the timing of migration. Humpback whale sightings significantly increased over the study period, but blue and gray whale counts did not, though there was variability across the time series. Date of breeding migration (departure) for all species showed little to no change, whereas date of migration towards feeding areas (arrival) occurred earlier for humpback and blue whales. Timing was significantly influenced by a mix of local oceanography, regional, and basin-scale climate variables. Earlier arrival time without concomitant earlier departure time results in longer periods when blue and humpback whales are at risk of entanglement in the Gulf of the Farallones. We maintain that these changes have increased whale exposure to pot and trap fishery gear off the central California coast during the spring, elevating the risk of entanglements. Humpback entanglement rates were significantly associated with increased counts and early arrival in central California. Actions to decrease the temporal overlap between whales and pot/trap fishing gear, particularly when whales arrive earlier in warm water years, would likely decrease the risk of entanglements.

Multiple pygmy blue whale acoustic populations in the Indian Ocean: whale song identifies a possible new population.

Blue whales were brought to the edge of extinction by commercial whaling in the twentieth century and their recovery rate in the Southern Hemisphere has been slow; they remain endangered. Blue whales, although the largest animals on Earth, are difficult to study in the Southern Hemisphere, thus their population structure, distribution and migration remain poorly known. Fortunately, blue whales produce powerful and stereotyped songs, which prove an effective clue for monitoring their different 'acoustic populations.' The DGD-Chagos song has been previously reported in the central Indian Ocean. A comparison of this song with the pygmy blue and Omura's whale songs shows that the Chagos song are likely produced by a distinct previously unknown pygmy blue whale population. These songs are a large part of the underwater soundscape in the tropical Indian Ocean and have been so for nearly two decades. Seasonal differences in song detections among our six recording sites suggest that the Chagos whales migrate from the eastern to western central Indian Ocean, around the Chagos Archipelago, then further east, up to the north of Western Australia, and possibly further north, as far as Sri Lanka. The Indian Ocean holds a greater diversity of blue whale populations than thought previously.

An open access dataset for developing automated detectors of Antarctic baleen whale sounds and performance evaluation of two commonly used detectors.

Since 2001, hundreds of thousands of hours of underwater acoustic recordings have been made throughout the Southern Ocean south of 60° S. Detailed analysis of the occurrence of marine mammal sounds in these circumpolar recordings could provide novel insights into their ecology, but manual inspection of the entirety of all recordings would be prohibitively time consuming and expensive. Automated signal processing methods have now developed to the point that they can be applied to these data in a cost-effective manner. However training and evaluating the efficacy of these automated signal processing methods still requires a representative annotated library of sounds to identify the true presence and absence of different sound types. This work presents such a library of annotated recordings for the purpose of training and evaluating automated detectors of Antarctic blue and fin whale calls. Creation of the library has focused on the annotation of a representative sample of recordings to ensure that automated algorithms can be developed and tested across a broad range of instruments, locations, environmental conditions, and years. To demonstrate the utility of the library, we characterise the performance of two automated detection algorithms that have been commonly used to detect stereotyped calls of blue and fin whales. The availability of this library will facilitate development of improved detectors for the acoustic presence of Southern Ocean blue and fin whales. It can also be expanded upon to facilitate standardization of subsequent analysis of spatiotemporal trends in call-density of these circumpolar species.

The largest of August Krogh animals: Physiology and biomechanics of the blue whale revisited.

The blue whale is the largest animal ever. This gigantism probably evolved to exploit seasonal krill blooms, where massive feasts allow for accumulation of large blubber reserves that can fuel their low mass specific metabolism during prolonged periods of fasting. Until recently, the physiology and biomechanics of blue whales could only be inferred from anatomical inspections, but the recent development of biologging tags now provide unique insights into how these ocean giants function and interact with their environment. Their mandibles, the largest bones ever to evolve, along with a highly expandable buccal cavity, enable an extreme and dynamic bulk feeding behavior. During a lunge feeding event, blue whales accelerate up to 5 m/s to engulf a volume prey laden water that is commensurate with the whale's gigantic body size. Perhaps due to the costs of such extreme foraging, their dive times of 10-15 min are much shorter than scaling would predict for their size. Like other diving animals, blue whales display a dive response with heart rates down to 4 BPM to prolong dive times and perhaps mitigate decompression sickness. Blue whales make the lowest and most energetic calls of any mammal with ocean traversing potential under natural ambient noise conditions. However, communication space may be severely reduced due to pervasive shipping noise. We hope that an increasing ability to study the physiology and behavior of blue whales and other marine megafauna will enable informed decisions and ensure our permanent co-existence in the face of increasing human encroachment into marine habitats.

Diel differences in blue whale (Balaenoptera musculus) dive behavior increase nighttime risk of ship strikes in northern Chilean Patagonia.

The northern Chilean Patagonia region is a key feeding ground and a nursing habitat in the southern hemisphere for blue whales (Balaenoptera musculus). From 2014 to 2019, during 6 separate research cruises, the dive behavior of 28 individual blue whales was investigated using bio-logging tags (DTAGs), generating ≈190 h of data. Whales dove to significantly greater depths during the day compared to nighttime (day: 32.6 ± 18.7 m; night: 6.2 ± 2.7 m; P < 0.01). During the night, most time was spent close to the surface (86% ± 9.4%; P < 0.01) and at depths of less than 12 m. From 2016 to 2019, active acoustics (scientific echosounders) were used to record prey (euphausiids) density and distribution simultaneously with whale diving data. Tagged whales appeared to perform dives relative to the vertical migration of prey during the day. The association between diurnal prey migration and shallow nighttime dive behavior suggests that blue whales are at increased risk of ship collisions during periods of darkness since the estimated maximum ship draft of vessels operating in the region is also ≈12 m. In recent decades, northern Chilean Patagonia has seen a large increase in marine traffic due to a boom in salmon aquaculture and the passenger ship industry. Vessel strike risks for large whales are likely underestimated in this region. Results reported in this study may be valuable for policy and mitigation decisions regarding conservation of the endangered blue whale.

Predicting suitable coastal habitat for sei whales, southern right whales and dolphins around the Falkland Islands.

Species distribution models (SDMs) are valuable tools for describing the occurrence of species and predicting suitable habitats. This study used generalized additive models (GAMs) and MaxEnt models to predict the relative densities of four cetacean species (sei whale Balaeanoptera borealis, southern right whale Eubalaena australis, Peale's dolphin Lagenorhynchus australis, and Commerson's dolphin Cephalorhynchus commersonii) in neritic waters (≤100 m depth) around the Falkland Islands, using boat survey data collected over three seasons (2017-2019). The model predictor variables (PVs) included remotely sensed environmental variables (sea surface temperature, SST, and chlorophyll-a concentration) and static geographical variables (e.g. water depth, distance to shore, slope). The GAM results explained 35 to 41% of the total deviance for sei whale, combined sei whales and unidentified large baleen whales, and Commerson's dolphins, but only 17% of the deviance for Peale's dolphins. The MaxEnt models for all species had low to moderate discriminatory power. The relative density of sei whales increased with SST in both models, and their predicted distribution was widespread across the inner shelf which is consistent with the use of Falklands' waters as a coastal summer feeding ground. Peale's dolphins and Commerson's dolphins were largely sympatric across the study area. However, the relative densities of Commerson's dolphins were generally predicted to be higher in nearshore, semi-enclosed, waters compared with Peale's dolphins, suggesting some habitat partitioning. The models for southern right whales performed poorly and the results were not considered meaningful, perhaps due to this species exhibiting fewer strong habitat preferences around the Falklands. The modelling results are applicable to marine spatial planning to identify where the occurrence of cetacean species and anthropogenic activities may most overlap. Additionally, the results can inform the process of delineating a potential Key Biodiversity Area for sei whales in the Falkland Islands.

Movements and behaviour of blue whales satellite tagged in an Australian upwelling system.

Knowledge about the movement ecology of endangered species is needed to identify biologically important areas and the spatio-temporal scale of potential human impacts on species. Blue whales (Balaenoptera musculus) are endangered due to twentieth century whaling and currently threatened by human activities. In Australia, they feed in the Great Southern Australian Coastal Upwelling System (GSACUS) during the austral summer. We investigate their movements, occupancy, behaviour, and environmental drivers to inform conservation management. Thirteen whales were satellite tagged, biopsy sampled and photo-identified in 2015. All were genetically confirmed to be of the pygmy subspecies (B. m. brevicauda). In the GSACUS, whales spent most of their time over the continental shelf and likely foraging in association with several seascape variables (sea surface temperature variability, depth, wind speed, sea surface height anomaly, and chlorophyll a). When whales left the region, they migrated west and then north along the Australian coast until they reached West Timor and Indonesia, where their movements indicated breeding or foraging behaviour. These results highlight the importance of the GSACUS as a foraging ground for pygmy blue whales inhabiting the eastern Indian Ocean and indicate the whales' migratory route to proposed breeding grounds off Indonesia. Information about the spatio-temporal scale of potential human impacts can now be used to protect this little-known subspecies of blue whale.

Blue whales.

Jeremy Goldbogen introduces blue whales, the largest animals to ever inhabit earth.

Animal-Borne Metrics Enable Acoustic Detection of Blue Whale Migration.

Linking individual and population scales is fundamental to many concepts in ecology [1], including migration [2, 3]. This behavior is a critical [4] yet increasingly threatened [5] part of the life history of diverse organisms. Research on migratory behavior is constrained by observational scale [2], limiting ecological understanding and precise management of migratory populations in expansive, inaccessible marine ecosystems [6]. This knowledge gap is magnified for dispersed oceanic predators such as endangered blue whales (Balaenoptera musculus). As capital breeders, blue whales migrate vast distances annually between foraging and breeding grounds, and their population fitness depends on synchrony of migration with phenology of prey populations [7, 8]. Despite previous studies of individual-level blue whale vocal behavior via bio-logging [9, 10] and population-level acoustic presence via passive acoustic monitoring [11], detection of the life history transition from foraging to migration remains challenging. Here, we integrate direct high-resolution measures of individual behavior and continuous broad-scale acoustic monitoring of regional song production (Figure 1A) to identify an acoustic signature of the transition from foraging to migration in the Northeast Pacific population. We find that foraging blue whales sing primarily at night, whereas migratory whales sing primarily during the day. The ability to acoustically detect population-level transitions in behavior provides a tool to more comprehensively study the life history, fitness, and plasticity of population behavior in a dispersed, capital breeding population. Real-time detection of this behavioral signal can also inform dynamic management efforts [12] to mitigate anthropogenic threats to this endangered population [13, 14]).

Modeling predator and prey hotspots: Management implications of baleen whale co-occurrence with krill in Central California.

As global ocean-bound commerce increases, managing human activities has become important in reducing conflict with threatened wildlife. This study investigates environmental factors determining abundance and distribution of blue whales (Balaenoptera musculus), humpback whales (Megaptera novaeangliae) and their prey (Euphausia pacifica and Thysanoessa spinifera) in central California. We provide insights into environmental drivers of the ecology and distribution of these species, model whale distributions and determine coincident hotspots of whales and their prey that will help decrease human threats to whales and protect critical feeding habitat. We developed separate predictive models of whale abundances (using negative binomial regression on count data) and krill abundance (using a two-part hurdlemodel combining logistic and negative binomial regressions) over a 14 year period (2004-2017). Variables included in situ surface and midwater oceanographic measures (temperature, salinity, and fluorescence), basin-scale climate indices, and bathymetric- and distance-related data. Predictions were applied to 1 km2 cells spanning the study area for May, June, July, and September during each of the 14 years of surveys to identify persistent distribution patterns. Both whales and krill were found to consistently use the northeast region of Cordell Bank, the Farallon Escarpment, and the shelf-break waters. The main identified blue whale hotspots were also krill hotspots, while co-occurrence was more limited and varied seasonally for humpback whales and krill. These results are valuable in identifying patterns in important areas of ecological interaction to assist management of whales. Areas north of Cordell Bank are of particular management concern since they overlap with the end of the San Francisco Bay northern shipping lane. Our findings can help decrease threats to whales, particularly in important foraging areas, by supporting implementation of vessel management and informing potential conflicts with other human uses.

Inter-annual decrease in pulse rate and peak frequency of Southeast Pacific blue whale song types.

A decrease in the frequency of two southeast Pacific blue whale song types was examined over decades, using acoustic data from several different sources in the eastern Pacific Ocean ranging between the Equator and Chilean Patagonia. The pulse rate of the song units as well as their peak frequency were measured using two different methods (summed auto-correlation and Fourier transform). The sources of error associated with each measurement were assessed. There was a linear decline in both parameters for the more common song type (southeast Pacific song type n.2) between 1997 to 2017. An abbreviated analysis, also showed a frequency decline in the scarcer southeast Pacific song type n.1 between 1970 to 2014, revealing that both song types are declining at similar rates. We discussed the use of measuring both pulse rate and peak frequency to examine the frequency decline. Finally, a comparison of the rates of frequency decline with other song types reported in the literature and a discussion on the reasons of the frequency shift are presented.

Timing is everything: Drivers of interannual variability in blue whale migration.

Blue whales need to time their migration from their breeding grounds to their feeding grounds to avoid missing peak prey abundances, but the cues they use for this are unknown. We examine migration timing (inferred from the local onset and cessation of blue whale calls recorded on seafloor-mounted hydrophones), environmental conditions (e.g., sea surface temperature anomalies and chlorophyll a), and prey (spring krill biomass from annual net tow surveys) during a 10 year period (2008-2017) in waters of the Southern California Region where blue whales feed in the summer. Colder sea surface temperature anomalies the previous season were correlated with greater krill biomass the following year, and earlier arrival by blue whales. Our results demonstrate a plastic response of blue whales to interannual variability and the importance of krill as a driving force behind migration timing. A decadal-scale increase in temperature due to climate change has led to blue whales extending their overall time in Southern California. By the end of our 10-year study, whales were arriving at the feeding grounds more than one month earlier, while their departure date did not change. Conservation strategies will need to account for increased anthropogenic threats resulting from longer times at the feeding grounds.

Baleen whale acoustic presence and behaviour at a Mid-Atlantic migratory habitat, the Azores Archipelago.

The identification of important areas during the annual life cycle of migratory animals, such as baleen whales, is vital for their conservation. In boreal springtime, fin and blue whales feed in the Azores on their way to northern latitudes while sei whales migrate through the archipelago with only occasional feeding. Little is known about their autumn or winter presence or their acoustic behaviour in temperate migratory habitats. This study used a 5-year acoustic data set collected by autonomous recorders in the Azores that were processed and analysed using an automated call detection and classification system. Fin and blue whales were acoustically present in the archipelago from autumn to spring with marked seasonal differences in the use of different call types. Diel patterns of calling activity were only found for fin whales with more calls during the day than night. Sei whales showed a bimodal distribution of acoustic presence in spring and autumn, corresponding to their expected migration patterns. Diel differences in sei whale calling varied with season and location. This work highlights the importance of the Azores as a migratory and wintering habitat for three species of baleen whales and provides novel information on their acoustic behaviour in a mid-Atlantic region.

Extreme bradycardia and tachycardia in the world's largest animal.

The biology of the blue whale has long fascinated physiologists because of the animal's extreme size. Despite high energetic demands from a large body, low mass-specific metabolic rates are likely powered by low heart rates. Diving bradycardia should slow blood oxygen depletion and enhance dive time available for foraging at depth. However, blue whales exhibit a high-cost feeding mechanism, lunge feeding, whereby large volumes of prey-laden water are intermittently engulfed and filtered during dives. This paradox of such a large, slowly beating heart and the high cost of lunge feeding represents a unique test of our understanding of cardiac function, hemodynamics, and physiological limits to body size. Here, we used an electrocardiogram (ECG)-depth recorder tag to measure blue whale heart rates during foraging dives as deep as 184 m and as long as 16.5 min. Heart rates during dives were typically 4 to 8 beats min-1 (bpm) and as low as 2 bpm, while after-dive surface heart rates were 25 to 37 bpm, near the estimated maximum heart rate possible. Despite extreme bradycardia, we recorded a 2.5-fold increase above diving heart rate minima during the powered ascent phase of feeding lunges followed by a gradual decrease of heart rate during the prolonged glide as engulfed water is filtered. These heart rate dynamics explain the unique hemodynamic design in rorqual whales consisting of a large-diameter, highly compliant, elastic aortic arch that allows the aorta to accommodate blood ejected by the heart and maintain blood flow during the long and variable pauses between heartbeats.

Using remote sensing to detect whale strandings in remote areas: The case of sei whales mass mortality in Chilean Patagonia.

We test the ability of Very High Resolution satellite (VHR) imagery to detect stranded whales using both manual and automated methods. We use the 2015 mass mortality event in the Gulf of Penas locality, central Patagonia, Chile, as an initial case study. This event was the largest known mass mortality of baleen whales, with at least 343 whales, mainly sei whales (Balaenoptera borealis), documented as stranding. However, even with such a large number of whales, due to the remote location of the gulf the strandings went unrecorded for several weeks. Aerial and boat surveys of the area were conducted two to four months after the mortality event. In this study we use 50cm resolution WorldView2 imagery to identify and count strandings from two archival images acquired just after the stranding event and two months before the aerial and ground surveys, and to test manual and automated methods of detecting stranded whales. Our findings show that whales are easily detected manually in the images but due to the heterogeneous colouration of decomposing whales, spectral indices are unsuitable for automatic detection. Our satellite counts suggest that, at the time the satellite images were taken, more whales were stranded than recorded in the aerial survey, possibly due to the non-comprehensive coverage of the aerial survey or movement of the carcases between survey acquisition. With even higher resolution imagery now available, satellite imagery may be a cost effective alternative to aerial surveys for future assessment of the extent of mass whale stranding events, especially in remote and inaccessible areas.