Fin whale song characteristics and potential subpopulation identity in the New York Bight.

Fin whale (Balaenoptera physalus) song can follow a highly consistent pattern, and regional differences in song patterns can be a valuable indicator of subpopulation identity and distribution. In the Northwest Atlantic, endangered fin whales are currently managed as a single stock despite previous identification of different regional song patterns, which indicates potential subpopulation structuring and vulnerability to anthropogenic disturbance if not managed accordingly. Here we document fin whale song in the New York Bight (NYB) from 2017 to 2020 using passive acoustic data to identify monthly and yearly trends in song patterns and to explore potential subpopulation structuring. The predominant song pattern observed was highly consistent with the pattern documented almost a decade prior in the NYB, with short inter-note intervals (INI) from fall-winter and long-INIs in the spring. However, in one song year the majority of songs were composed of long-INIs. This change in song pattern could be due to a shift in fin whale behavior or possibly multiple fin whale subpopulations using the NYB. Fin whales in the NYB may be particularly vulnerable to disturbance given the increasing anthropogenic pressures in this region, and further research into subpopulation structuring is needed to ensure adequate management of these endangered whales.

Acoustic detection range of right whale upcalls identified in near-real time from a moored buoy and a Slocum glider.

The goal of this study was to characterize the detection range of a near real-time baleen whale detection system, the digital acoustic monitoring instrument/low-frequency detection and classification system (DMON/LFDCS), equipped on a Slocum glider and a moored buoy. As a reference, a hydrophone array was deployed alongside the glider and buoy at a shallow-water site southwest of Martha's Vineyard (Massachusetts, USA) over a four-week period in spring 2017. A call-by-call comparison between North Atlantic right whale upcalls localized with the array (n = 541) and those detected by the glider or buoy was used to estimate the detection function for each DMON/LFDCS platform. The probability of detection was influenced by range, ambient noise level, platform depth, detection process, review protocol, and calling rate. The conservative analysis of near real-time pitch tracks suggested that, under typical conditions, a 0.33 probability of detection of a single call occurred at 6.2 km for the buoy and 8.6-13.4 km for the glider (depending on glider depth), while a 0.10 probability of detection of a single call occurred at 14.4 m for the buoy and 22.6-27.5 km for the glider. Probability of detection is predicted to increase substantially at all ranges if more than one call is available for detection.

Near real-time detection of low-frequency baleen whale calls from an autonomous surface vehicle: Implementation, evaluation, and remaining challenges.

Mitigation of threats posed to marine mammals by human activities can be greatly improved with a better understanding of animal occurrence in real time. Recent advancements have enabled low-power passive acoustic systems to be integrated into long-endurance autonomous platforms for persistent near real-time monitoring of marine mammals via the sounds they produce. Here, the integration of a passive acoustic instrument capable of real-time detection and classification of low-frequency (LF) tonal sounds with a Liquid Robotics wave glider is reported. The goal of the integration was to enable monitoring of LF calls produced by baleen whales over periods of several months. Mechanical noises produced by the platform were significantly reduced by lubricating moving parts with polytetrafluoroethylene, incorporating rubber and springs to decelerate moving parts and shock mounting hydrophones. Flow noise was reduced with the development of a 21-element hydrophone array. Surface noise produced by breaking waves was not mitigated despite experimentation with baffles. Compared to a well-characterized moored passive acoustic monitoring buoy, the system greatly underestimated the occurrence of sei, fin, and North Atlantic right whales during a 37-d deployment, and therefore is not suitable in its current configuration for use in scientific or management applications for these species at this time.

Self-localization of buoyless fishing gear and other objects on the sea floor.

End lines used in commercial trap/pot fishing pose a significant entanglement risk to whales, sea turtles, and sharks. Removal of these ropes for buoyless fishing is being considered by the United States and Canadian governments, but a method to systematically locate the gear without an attached buoy is required. A method was developed for an acoustic modem to self-localize and broadcast its location to nearby ships to minimize gear conflict, optimize power consumption, and reduce lost gear. This method was implemented using a research modem that self-localized to within 5 m of its estimated location on the sea floor.

Bowhead whales use two foraging strategies in response to fine-scale differences in zooplankton vertical distribution.

As zooplanktivorous predators, bowhead whales (Balaena mysticetus) must routinely locate patches of prey that are energy-rich enough to meet their metabolic needs. However, little is known about how the quality and quantity of prey might influence their feeding behaviours. We addressed this question using a new approach that included: (1) multi-scale biologging and unmanned aerial system observations of bowhead whales in Cumberland Sound, Nunavut (Canada), and (2) an optical plankton counter (OPC) and net collections to identify and enumerate copepod prey species through the water column. The OPC data revealed two prey layers comprised almost exclusively of lipid-rich calanoid copepods. The deep layer contained fewer, but larger, particles (10% greater overall biomass) than the shallow prey layer. Dive data indicated that the whales conducted long deep Square-shaped dives (80% of dives; averaging depth of 260.4 m) and short shallow Square-shaped dives (16%; averaging depth of 22.5 m) to feed. The whales tended to dive proportionally more to the greater biomass of zooplankton that occurred at depth. Combining behavioural recordings with prey sampling showed a more complex feeding ecology than previously understood, and provides a means to evaluate the energetic balance of individuals under current environmental conditions.

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.

Inter-individual variability in copepod microbiomes reveals bacterial networks linked to host physiology.

Copepods harbor diverse bacterial communities, which collectively carry out key biogeochemical transformations in the ocean. However, bulk copepod sampling averages over the variability in their associated bacterial communities, thereby limiting our understanding of the nature and specificity of copepod-bacteria associations. Here, we characterize the bacterial communities associated with nearly 200 individual Calanus finmarchicus copepods transitioning from active growth to diapause. We find that all individual copepods sampled share a small set of "core" operational taxonomic units (OTUs), a subset of which have also been found associated with other marine copepod species in different geographic locations. However, most OTUs are patchily distributed across individual copepods, thereby driving community differences across individuals. Among patchily distributed OTUs, we identified groups of OTUs correlated with common ecological drivers. For instance, a group of OTUs positively correlated with recent copepod feeding served to differentiate largely active growing copepods from those entering diapause. Together, our results underscore the power of individual-level sampling for understanding host-microbiome relationships.

Correction: Evidence of molting and the function of "rock-nosing" behavior in bowhead whales in the eastern Canadian Arctic.

[This corrects the article DOI: 10.1371/journal.pone.0186156.].

Evidence of molting and the function of "rock-nosing" behavior in bowhead whales in the eastern Canadian Arctic.

Bowhead whales (Balaena mysticetus) have a nearly circumpolar distribution, and occasionally occupy warmer shallow coastal areas during summertime that may facilitate molting. However, relatively little is known about the occurrence of molting and associated behaviors in bowhead whales. We opportunistically observed whales in Cumberland Sound, Nunavut, Canada with skin irregularities consistent with molting during August 2014, and collected a skin sample from a biopsied whale that revealed loose epidermis and sloughing. During August 2016, we flew a small unmanned aerial system (sUAS) over whales to take video and still images to: 1) determine unique individuals; 2) estimate the proportion of the body of unique individuals that exhibited sloughing skin; 3) determine the presence or absence of superficial lines representative of rock-rubbing behavior; and 4) measure body lengths to infer age-class. The still images revealed that all individuals (n = 81 whales) were sloughing skin, and that nearly 40% of them had mottled skin over more than two-thirds of their bodies. The video images captured bowhead whales rubbing on large rocks in shallow, coastal areas-likely to facilitate molting. Molting and rock rubbing appears to be pervasive during late summer for whales in the eastern Canadian Arctic.

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

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

Coping with copepods: do right whales (Eubalaena glacialis) forage visually in dark waters?

North Atlantic right whales (Eubalaena glacialis) feed during the spring and early summer in marine waters off the northeast coast of North America. Their food primarily consists of planktonic copepods, Calanus finmarchicus, which they consume in large numbers by ram filter feeding. The coastal waters where these whales forage are turbid, but they successfully locate copepod swarms during the day at depths exceeding 100 m, where light is very dim and copepod patches may be difficult to see. Using models of E. glacialis visual sensitivity together with measurements of light in waters near Cape Cod where they feed and of light attenuation by living copepods in seawater, we evaluated the potential for visual foraging by these whales. Our results suggest that vision may be useful for finding copepod patches, particularly if E. glacialis searches overhead for silhouetted masses or layers of copepods. This should permit the whales to locate C. finmarchicus visually throughout most daylight hours at depths throughout their foraging range. Looking laterally, the whales might also be able to see copepod patches at short range near the surface.This article is part of the themed issue 'Vision in dim light'.

The Physiology and Ecology of Diapause in Marine Copepods.

Diapause is a type of dormancy that requires preparation, typically precedes the onset of unfavorable conditions, and necessitates a period of arrest before development can proceed. Two ecologically important groups of copepods have incorporated diapausing stages into their life histories. In freshwater, estuarine, and coastal environments, species within the Centropagoidea superfamily can produce resting eggs containing embryos that may be quiescent, diapausing, or in some intermediate state. Resting eggs sink into the sediments, remain viable over months to years, and form a reservoir from which the planktonic population is reestablished. In coastal and oceanic environments, copepods within the Calanidae and Eucalanidae families can enter diapause during late juvenile (copepodid) or adult stages. These copepods accumulate large amounts of lipids before they migrate into deep water and diapause for several months. Through respiration, diapausing copepods may sequester more carbon in the deep ocean than any other biogeochemical process, and changes in diapause phenology associated with climate change (particularly reduction in diapause duration) could have a significant impact not only on regional ecosystems, but on global climate as well.

Transcriptional Profiling of Metabolic Transitions during Development and Diapause Preparation in the Copepod Calanus finmarchicus.

Calanus finmarchicus, like many other copepods in the family Calanidae, can enter into a facultative diapause during the last juvenile phase (fifth copepodid, C5) to enable survival during unfavorable periods. Diapause is essential to the persistence of Calanus populations and profoundly impacts energy flow within oceanic ecosystems, yet regulation of diapause is not understood in these animals. Transcriptional profiling has begun to provide insight into metabolic changes occurring as C. finmarchicus prepares for and enters into diapause or skips diapause to prepare for the terminal molt. In particular, components of the glycolysis, pentose phosphate and lipid synthesis pathways are upregulated early in the C5 stage when lipid stores are low. Currently, our ability to identify metabolic patterns is limited by the incomplete functional annotation of the C. finmarchicus transcriptome. Such limitations are widespread among studies of non-model organisms and addressing them should be a priority for future research. In addition, integrating the results across multiple emerging complementary transcriptomic studies will provide a more complete picture of copepod physiology than isolated studies. Ultimately, identifying molecular markers of copepod physiology could enable robust identification of animals preparing to enter into diapause and ultimately lead to a greatly improved understanding of diapause regulation.

Assessing performance of Bayesian state-space models fit to Argos satellite telemetry locations processed with Kalman filtering.

Argos recently implemented a new algorithm to calculate locations of satellite-tracked animals that uses a Kalman filter (KF). The KF algorithm is reported to increase the number and accuracy of estimated positions over the traditional Least Squares (LS) algorithm, with potential advantages to the application of state-space methods to model animal movement data. We tested the performance of two Bayesian state-space models (SSMs) fitted to satellite tracking data processed with KF algorithm. Tracks from 7 harbour seals (Phoca vitulina) tagged with ARGOS satellite transmitters equipped with Fastloc GPS loggers were used to calculate the error of locations estimated from SSMs fitted to KF and LS data, by comparing those to "true" GPS locations. Data on 6 fin whales (Balaenoptera physalus) were used to investigate consistency in movement parameters, location and behavioural states estimated by switching state-space models (SSSM) fitted to data derived from KF and LS methods. The model fit to KF locations improved the accuracy of seal trips by 27% over the LS model. 82% of locations predicted from the KF model and 73% of locations from the LS model were <5 km from the corresponding interpolated GPS position. Uncertainty in KF model estimates (5.6 ± 5.6 km) was nearly half that of LS estimates (11.6 ± 8.4 km). Accuracy of KF and LS modelled locations was sensitive to precision but not to observation frequency or temporal resolution of raw Argos data. On average, 88% of whale locations estimated by KF models fell within the 95% probability ellipse of paired locations from LS models. Precision of KF locations for whales was generally higher. Whales' behavioural mode inferred by KF models matched the classification from LS models in 94% of the cases. State-space models fit to KF data can improve spatial accuracy of location estimates over LS models and produce equally reliable behavioural estimates.

Transcriptional profiling of reproductive development, lipid storage and molting throughout the last juvenile stage of the marine copepod Calanus finmarchicus.

INTRODUCTION: Calanus finmarchicus, a highly abundant copepod that is an important primary consumer in North Atlantic ecosystems, has a flexible life history in which copepods in the last juvenile developmental stage (fifth copepodid, C5) may either delay maturation and enter diapause or molt directly into adults. The factors that regulate this developmental plasticity are poorly understood, and few tools have been developed to assess the physiological condition of individual copepods. RESULTS: We sampled a cultured population of C. finmarchicus copepods daily throughout the C5 stage and assessed molt stage progression, gonad development and lipid storage. We used high-throughput sequencing to identify genes that were differentially expressed during progression through the molt stage and then used qPCR to profile daily expression of individual genes. Based on expression profiles of twelve genes, samples were statistically clustered into three groups: (1) an early period occurring prior to separation of the cuticle from the epidermis (apolysis) when expression of genes associated with lipid synthesis and transport (FABP and ELOV) and two nuclear receptors (ERR and HR78) was highest, (2) a middle period of rapid change in both gene expression and physiological condition, including local minima and maxima in several nuclear receptors (FTZ-F1, HR38b, and EcR), and (3) a late period when gonads were differentiated and expression of genes associated with molting (Torso-like, HR38a) peaked. The ratio of Torso-like to HR38b strongly differentiated the early and late groups. CONCLUSIONS: This study provides the first dynamic profiles of gene expression anchored with morphological markers of lipid accumulation, development and gonad maturation throughout a copepod molt cycle. Transcriptomic profiling revealed significant changes over the molt cycle in genes with presumed roles in lipid synthesis, molt regulation and gonad development, suggestive of a coupling of these processes in Calanus finmarchicus. Finally, we identified gene expression profiles that strongly differentiate between early and late development within the C5 copepodid stage. We anticipate that these findings and continued development of robust gene expression biomarkers that distinguish between diapause preparation and continuous development will ultimately enable novel studies of the intrinsic and extrinsic factors that govern diapause initiation in Calanus finmarchicus.

North Atlantic blue and fin whales suspend their spring migration to forage in middle latitudes: building up energy reserves for the journey?

The need to balance energy reserves during migration is a critical factor for most long-distance migrants and an important determinant of migratory strategies in birds, insects and land mammals. Large baleen whales migrate annually between foraging and breeding sites, crossing vast ocean areas where food is seldom abundant. How whales respond to the demands and constraints of such long migrations remains unknown. We applied a behaviour discriminating hierarchical state-space model to the satellite tracking data of 12 fin whales and 3 blue whales tagged off the Azores, to investigate their movements, behaviour (transiting and area-restricted search, ARS) and daily activity cycles during the spring migration. Fin and blue whales remained at middle latitudes for prolonged periods, spending most of their time there in ARS behaviour. While near the Azores, fin whale ARS behaviour occurred within a restricted area, with a high degree of overlap among whales. There were noticeable behavioural differences along the migratory pathway of fin whales tracked to higher latitudes: ARS occurred only in the Azores and north of 56°N, whereas in between these areas whales travelled at higher overall speeds while maintaining a nearly direct trajectory. This suggests fin whales may alternate periods of active migration with periods of extended use of specific habitats along the migratory route. ARS behaviour in blue whales occurred over a much wider area as whales slowly progressed northwards. The tracks of these whales terminated still at middle latitudes, before any behavioural switch was detected. Fin whales exhibited behavioural-specific diel rhythms in swimming speed but these varied significantly between geographic areas, possibly due to differences in the day-night cycle across areas. Finally, we show a link between fin whales seen in the Azores and those summering in eastern Greenland-western Iceland along a migratory corridor located in central Atlantic waters.

Real-time reporting of baleen whale passive acoustic detections from ocean gliders.

In the past decade, much progress has been made in real-time passive acoustic monitoring of marine mammal occurrence and distribution from autonomous platforms (e.g., gliders, floats, buoys), but current systems focus primarily on a single call type produced by a single species, often from a single location. A hardware and software system was developed to detect, classify, and report 14 call types produced by 4 species of baleen whales in real time from ocean gliders. During a 3-week deployment in the central Gulf of Maine in late November and early December 2012, two gliders reported over 25,000 acoustic detections attributed to fin, humpback, sei, and right whales. The overall false detection rate for individual calls was 14%, and for right, humpback, and fin whales, false predictions of occurrence during 15-min reporting periods were 5% or less. Transmitted pitch tracks--compact representations of sounds--allowed unambiguous identification of both humpback and fin whale song. Of the ten cases when whales were sighted during aerial or shipboard surveys and a glider was within 20 km of the sighting location, nine were accompanied by real-time acoustic detections of the same species by the glider within ±12 h of the sighting time.

Long distance passive localization of vocalizing sei whales using an acoustic normal mode approach.

During a 2 day period in mid-September 2006, more than 200, unconfirmed but identifiable, sei whale (Balaenoptera borealis) calls were collected as incidental data during a multidisciplinary oceanography and acoustics experiment on the shelf off New Jersey. Using a combined vertical and horizontal acoustic receiving array, sei whale movements were tracked over long distances (up to tens of kilometers) using a normal mode back propagation technique. This approach uses low-frequency, broadband passive sei whale call receptions from a single-station, two-dimensional hydrophone array to perform long distance localization and tracking by exploiting the dispersive nature of propagating normal modes in a shallow water environment. The back propagation approach is examined for accuracy and application to tracking the sei whale vocalizations identified in the vertical and horizontal array signals. This passive whale tracking, combined with the intensive oceanography measurements performed during the experiment, was also used to examine sei whale movements in relation to oceanographic features observed in this region.

A generalized baleen whale call detection and classification system.

Passive acoustic monitoring allows the assessment of marine mammal occurrence and distribution at greater temporal and spatial scales than is now possible with traditional visual surveys. However, the large volume of acoustic data and the lengthy and laborious task of manually analyzing these data have hindered broad application of this technique. To overcome these limitations, a generalized automated detection and classification system (DCS) was developed to efficiently and accurately identify low-frequency baleen whale calls. The DCS (1) accounts for persistent narrowband and transient broadband noise, (2) characterizes temporal variation of dominant call frequencies via pitch-tracking, and (3) classifies calls based on attributes of the resulting pitch tracks using quadratic discriminant function analysis (QDFA). Automated detections of sei whale (Balaenoptera borealis) downsweep calls and North Atlantic right whale (Eubalaena glacialis) upcalls were evaluated using recordings collected in the southwestern Gulf of Maine during the spring seasons of 2006 and 2007. The accuracy of the DCS was similar to that of a human analyst: variability in differences between the DCS and an analyst was similar to that between independent analysts, and temporal variability in call rates was similar among the DCS and several analysts.

Heat shock protein expression during stress and diapause in the marine copepod Calanus finmarchicus.

Calanoid copepods, such as Calanus finmarchicus, are a key component of marine food webs. C. finmarchicus undergo a facultative diapause during juvenile development, which profoundly affects their seasonal distribution and availability to their predators. The current ignorance of how copepod diapause is regulated limits understanding of copepod population dynamics, distribution, and ecosystem interactions. Heat shock proteins (Hsps) are a superfamily of molecular chaperones characteristically upregulated in response to stress conditions and frequently associated with diapause in other taxa. In this study, 8 heat shock proteins were identified in C. finmarchicus C5 copepodids (Hsp21, Hsp22, p26, Hsp90, and 4 forms of Hsp70), and expression of these transcripts was characterized in response to handling stress and in association with diapause. Hsp21, Hsp22, and Hsp70A (cytosolic subfamily) were induced by handling stress. Expression of Hsp70A was also elevated in shallow active copepodids relative to deep diapausing copepodids, which may reflect induction of this gene by varied stressors in active animals. In contrast, expression of Hsp22 was elevated in deep diapausing animals; Hsp22 may play a role both in short-term stress responses and in protecting proteins from degradation during diapause. Expression of most of the Hsps examined did not vary in response to diapause, perhaps because the diapause of C. finmarchicus is not associated with the extreme environmental conditions (e.g., freezing and desiccation) experienced by many other taxa, such as overwintering insects or Artemia cysts.