Search

Short-finned pilot whales (Globicephala macrorhynchus) are large, deep-diving predators with diverse foraging strategies, but little is known about their echolocation. To quantify the source properties of short-finned pilot whale clicks, we made 15 deployments off the coast of Tenerife of a deep-water hydrophone array consisting of seven autonomous time-synced hydrophone recorders (SoundTraps), enabling acoustic localization and quantification of click source parameters. Of 8185 recorded pilot whale clicks, 47 were classified as being recorded on-axis, with a mean peak-to-peak source level (SL) of 181 ± 7 dB re 1 μPa, a centroid frequency of 40 ± 4 kHz, and a duration of 57 ± 23 μs. A fit to a piston model yielded an estimated half-power (-3 dB) beam width of 13.7° [95% confidence interval (CI) 13.2°-14.5°] and a mean directivity index (DI) of 22.6 dB (95% CI 22.5-22.9 dB). These measured SLs and DIs are surprisingly low for a deep-diving toothed whale, suggesting we sampled the short-finned pilot whales in a context with little need for operating a long-range biosonar. The substantial spectral overlap with beaked whale clicks emitted in similar deep-water habitats implies that pilot whale clicks may constitute a common source of false detections in beaked whale passive acoustic monitoring efforts.

Information on the gut microbiome composition of different mammals could provide novel insights into the evolution of mammals and succession of microbial communities in different hosts. However, there is limited information on the gut microbiome composition of marine mammals, especially cetaceans, because of sampling constraints. In this study, we investigated the diversity and composition of microbial communities in the stomach, midgut, and hindgut of two stranded short-finned pilot whales (Globicephala macrorhynchus) and hindgut of a stranded pygmy sperm whale (Kogia breviceps) by using 16S rRNA gene amplicon sequencing technology. On the basis of the 50 most abundant operational taxonomic units, principal coordinate analysis, and non-metric multidimensional scaling analysis, we confirmed that the gut microbial communities of the three whales were different. Our results revealed that the gut microbiome of one stranded short-finned pilot whale GM16 was dominated by Firmicutes (mainly Clostridium) and Fusobacteria; whereas that of the other pilot whale GM19 was composed of Gammaproteobacteria and Bacteroidetes (mainly Vibrio and Bacteroides, respectively), probably caused by intestinal disease and antibiotic treatment. The gut microbiome of the pygmy sperm whale was dominated by Firmicutes and Bacteroidetes. Moreover, different gastrointestinal tract regions harbored different microbial community structures. To our knowledge, this is the first report of the gut microbiome of short-finned pilot whales, and our findings will expand our current knowledge on microbial diversity and composition in the gastrointestinal tract of cetaceans. This article is protected by copyright. All rights reserved.

The visual system of cetaceans is at best poorly understood. With a handful of electrophysiological studies and a limited number of histological preparations from well-preserved specimen, the investigation of the principles underlying the cortical organization in cetaceans remains a challenge. In the course of our current investigation, we identified the transition from V2 to V1 in the long-finned pilot whale Globicephala melas, only recognizable through immunocytochemistry, and a similar if not homologue transition in the sheep Ovis aries. Our results emphasize the importance of differential pattern recognition in which the application of different markers uncovers a diversity in a delphinid's cortex, formerly widely considered as uniform and archetypal. In fact, the evidence that we present suggests the existence of relatively unacknowledged areas beyond the well-known sensory territories in cetaceans.

Summary Polysaccharide storage myopathies have been described in several animal species and are characterized by periodic acid–Schiff (PAS)-positive, diastase-resistant intrasarcoplasmic inclusions in myocytes. Skeletal and cardiac muscle samples from a subset of a single pod of stranded long-finned pilot whales (Globicephala melas) were evaluated by light and transmission electron microscopy. Twelve individuals demonstrated sporadic basophilic packets of PAS-positive, diastase-resistant complex polysaccharide material, either centrally or peripherally, in skeletal and cardiac myocytes. Few microscopic myopathic changes were found but included focal inflammation and internalized nuclei. Ultrastructurally, the inclusions consisted of loosely arranged, tangled filaments and were not membrane-bound, which is consistent with polyglucosan inclusions. Within skeletal muscle, the number of inclusions had a marginal statistically significant (P = 0.0536) correlation with length, as a proxy for age, suggesting that such inclusions in skeletal muscles may be age-related, although the cause remains unclear.

While in the northern hemisphere, many studies have been conducted on the vocal repertoire of long-finned pilot whales (Globicephala melas), no such study has been conducted in the southern hemisphere. Presented here, is the first study on the vocalisations of long-finned pilot whales along the southern coast of mainland Australia. Multiple measures were taken of 2028 vocalisations recorded over five years in several locations. These vocalisations included tonal sounds with and without overtones, sounds of burst-pulse character, graded sounds, biphonations, and calls of multiple components. Vocalisations were further categorised based on spectrographic features into 18 contour classes. Altogether, vocalisations ranged from approximately 200 Hz to 25 kHz in fundamental frequency and from 0.03 s to 2.07 s in duration. These measures compared well with those from northern hemisphere pilot whales. Some call types were almost identical to northern hemisphere vocalisations, even though the geographic ranges of the two populations are far apart. Other call types were unique to Australia. Striking similarities with calls of short-finned pilot whales (Globicephala macrorhynchus) and sometimes sympatric killer whales (Orcinus orca) were also found. Theories for call convergence and divergence are discussed.

Ecological data on marine mammal parasites represent an excellent opportunity to expand our understanding of host-parasite systems. In this study, we used a dataset of intestinal helminth parasites on 167 long-finned pilot whales Globicephala melas (Traill, 1809) from seven localities in the Faroe Islands to evaluate the extent to which the host's age and sex influence the occurrence, richness, and nested pattern of helminth parasites and the importance of individual hosts to the helminth community. We found positive effects of age on both the occurrence and richness of helminths. Older host individuals showed an ordered accumulation of parasites, as evidenced by the nested pattern in their composition. Males had a higher occurrence of parasites than females, but the richness of helminths did not differ between sexes. Our findings suggest that differences in host-parasite interactions in long-finned pilot whales result mainly from age-structured variations in biological and behavioral characteristics.

The number of social contacts of mammals is positively correlated with the diversity of their gut microbes. There is some evidence that sociality also affects microbes in the respiratory tract. We tested whether the airway microbiota of cetacean species differ depending on the whales' level of sociality. We sampled the blow of blue (Balaenoptera musculus), grey (Eschrichtius robustus), humpback (Megaptera novaeangliae) and long-finned pilot whales (PWs) (Globicephala melas) and analysed the blow microbiota by barcode tag sequencing targeting the V4 region of the bacterial 16S rRNA gene. Humpback whales (HWs) show higher levels of sociality than blue (BW) and grey (GW), while PWs are the most gregarious among the four species. The blow samples of the HWs showed the highest richness and diversity. HWs were also the only species with a species-specific clustering of their microbial community composition and a relatively large number of core taxa. Therefore, we conclude that it cannot be sociality alone shaping the diversity and composition of airway microbiota. We suggest the whale species' lung volume and size of the plume of exhaled air as an additional factor impacting the transmission potential of blow microbiota from one individual whale to another.

Nucleic acid-derived indices such as RNA/DNA ratios have been successfully applied as ecophysiological indicators to assess growth, nutritional condition and health status in marine organisms given that they provide a measure of tissue protein reserves, which is known to vary depending on changes in the environment. Yet, the use of these biochemical indices on highly mobile large predators is scarce. In this study, we tested the applicability of using nucleic acids to provide insights on the ecophysiological traits of two marine mammal species (common bottlenose dolphins and short-finned pilot whales) and explored potential related factors (species, sex, season, and residency pattern), using skin tissue (obtained from biopsy darts) of apparently healthy and adult free-ranging animals. Significantly higher RNA/DNA ratios were obtained for bottlenose dolphins (p

Although lipophilic compounds have been the focus of numerous studies in marine mammals, their association with lipids is widely accepted, but rarely scrutinized. This pilot study aimed to investigate potential relationships between individual lipids from different lipid classes identified through a non-targeted Nuclear Magnetic Resonance (NMR) based lipidomics approach and legacy POPs in the blubber of long-finned pilot whales, sperm whales, common bottlenose dolphins, and Indo-Pacific bottlenose dolphins. Concentrations of selected POPs such as HCB and HCHs in sperm whales from Tasmania were found to differ from those in long-finned pilot whales and common bottlenose dolphins from the same location. Profiles of NMR spectra measured in blubber of sperm whales were also distinctly different compared to the pilot whales and common bottlenose dolphins. Two groups of Indo-Pacific bottlenose dolphins from South Australia that were 20 years apart showed highly comparable profiles of NMR signals despite having higher concentrations of several POP classes in the more recent group. More specific correlations were investigated between selected POPs (n = 12) and all detected NMR signals (n = 63) in all species. Outcomes were species-specific, but difficult to interpret due to the lack of available literature for marine mammals and the small sample sizes per species. Because of the key role of lipids in the bioaccumulation of POPs and in the incidence of diseases, more attention should be given to the identification and characterization of lipid species in future toxicological studies. However, future studies should focus on one marine mammal species to increase sample sizes and limit the number of confounding factors, such as diet, that can influence POP and lipid levels and profiles.

The sound-transmission, beam-formation, and sound-reception processes of a short-finned pilot whale (Globicephala macrorhynchus) were investigated using computed tomography (CT) scanning and numerical simulation. The results showed that sound propagations in the forehead were modulated by the upper jaw, air components, and soft tissues, which attributed to the beam formation in the external acoustic field. These structures owned different acoustic impedance and formed a multiphasic sound transmission system that can modulate sounds into a beam. The reception pathways composed of the solid mandible and acoustic fats in the lower head conducted sounds into the tympano-periotic complex. In the simulations, sounds were emitted in the forehead transmission system and propagated into water to interrogate a steel cylinder. The resulting echoes can be interpreted from multiple perspectives, including amplitude, waveform, and spectrum, to obtain the acoustic cues of the steel cylinder. By taking the short-finned pilot whale as an example, this study provides meaningful information to further deepen our understanding of biosonar system operations, and may expand sound-reception theory in odontocetes.

Over the last decades global warming has caused an increase in ocean temperature, acidification and oxygen loss which has led to changes in nutrient cycling and primary production affecting marine species at multiple trophic levels. While knowledge about the impacts of climate change in cetacean's species is still scarce, practitioners and policymakers need information about the species at risk to guide the implementation of conservation measures. To assess cetacean's vulnerability to climate change in the biogeographic region of Macaronesia, we adapted the Marine Mammal Climate Vulnerability Assessment (MMCVA) method and applied it to 21 species management units using an expert elicitation approach. Results showed that over half (62%) of the units assessed presented Very High (5 units) or High (8 units) vulnerability scores. Very High vulnerability scores were found in archipelago associated units of short-finned pilot whales (Globicephala macrorhynchus) and common bottlenose dolphins (Tursiops truncatus), namely in the Canary Islands and Madeira, as well as Risso's dolphins (Grampus griseus) in the Canary Islands. Overall, certainty scores ranged from Very High to Moderate for 67% of units. Over 50% of units showed a high potential for distribution, abundance and phenology changes as a response to climate change. With this study we target current and future information needs of conservation managers in the region, and guide research and monitoring efforts, while contributing to the improvement and validation of trait-based vulnerability approaches under a changing climate. This research was supported by the European Union's Horizon 2020 research and innovation programme under grant agreement No 776661, project “SOCLIMPACT—DownScaling CLImate imPACTs and decarbonisation pathways in EU islands and enhancing socioeconomic and non-market evaluation of Climate Change for Europe, for 2050 and beyond”. info:eu-repo/semantics/publishedVersion

Vessel noise is a primary driver of behavioural disturbance in cetaceans, which are targeted during whale-watch activities. Despite the growing, global effort for implementing best-practice principles, to date, there are no regulations on whale-watch vessel noise levels. Here, we test the hypothesis that a whale-watch vessel with a low noise emission will not elicit short-term behavioural responses in toothed whales compared to a vessel with a louder engine. We measured behavioural responses (n = 36) of short-finned pilot whales (Globicephala macrorhynchus) to whale-watch vessel approaches (range 60 m, speed 1.5 kn). Treatment approaches with a quieter electric engine (136–140 dB) compared to the same vessel with a louder petrol engine (151–139 dB) (low-frequency–mid-frequency weighted source levels, re 1 µPa RMS @ 1 m) were examined. Focal whales were resting mother and calves in small group sizes. During petrol engine treatments, the mother’s mean resting time decreased by 29% compared to the control (GLM, p = 0.009). The mean proportion of time nursing for the calf was significantly influenced by petrol engine vessel passes, with a 81% decrease compared to the control (GLM, p = 0.01). There were no significant effects on behaviour from the quieter electric engine. Thus, to minimise disturbance on the activity budget of pilot whales, whale-watch vessels would ideally have source levels as low as possible, below 150 dB re 1 µPa RMS @ 1 m and perceived above ambient noise.

Marine metal pollution is an emerging concern for human, animal, and ecosystem health. We considered metal pollution in the Sea of Cortez, which is a relatively isolated sea rich in biodiversity. Here there are potentially significant anthropogenic inputs of pollution from agriculture and metal mining. We considered the levels of 23 heavy metals and selenium in seven distinct cetacean species found in the area. Our efforts considered two different periods of time: 1999 and 2016/17. We considered the metal levels in relation to (1) all species together across years, (2) differences between suborders Odontoceti and Mysticeti, (3) each species individually across years, and (4) gender differences for each of these comparisons. We further compared metal levels found in sperm whale skin samples collected during these voyages to a previous voyage in 1999, to assess changes in metal levels over a longer timescale. The metals Mg, Fe, Al, and Zn were found at the highest concentrations across all species and all years. For sperm whales, we observed decreased metal levels from 1999 to 2016/2017, except for iron (Fe), nickel (Ni), and chromium (Cr), which either increased or did not change during this time period. These results indicate a recent change in the metal input to the Sea of Cortez, which may indicate a decreased concern for human, animal, and ecosystem health for some metals, but raises concern for the genotoxic metals Cr and Ni. This work was supported by NIEHS grant ES016893 (J.P.W.) and numerous donors to the Wise Laboratory.

Fat-level measurements used to indicate individual body condition and fitness are useful only when taken at a region along the body where fat responds to variations in caloric intake. Investigations to identify appropriate species-specific regions are limited, especially for cetaceans that have a specialized fat (blubber) that serves as an energy reserve and provides insulation. Over 18 mo, body mass of six pilot whales varied (range: 50-172 kg), and although caloric intake increased when water temperatures were lower, generally the best-fitting state-space model for length-adjusted mass was based on a single factor, caloric intake. After correcting for body length (range: 330-447 cm), the slope for blubber thickness and "blubber ring" thickness (average blubber thickness along a girth) in relation to body mass was positive and had a

For most marine vertebrates, chemical cues provide crucial information during navigation and foraging, but their use by cetaceans is still poorly understood. In contrast to baleen whales, toothed whales (odontocetes) are scarcely equipped for chemoreception: they lack the conventional anatomical structures (i.e., olfactory epithelium, nerves and bulbs) involved in olfaction and have reduced taste buds on the tongue. Several behavioral studies have however shown that captive dolphins can perceive chemical solutions, including odorants, in their oral cavity. To investigate whether odontocetes could use infochemicals in their foraging ecology, we implemented a behavioral response experiment in wild bottlenose dolphins and long-finned pilot whales. We tested dimethyl sulfide (DMS) as a potentially attractive stimulus since it is a chemical signature of highly productive marine areas, known to attract several marine predators including fishes and seabirds. We assessed cetacean responses to DMS exposure by analyzing their movements and surface behaviors recorded by onboard observers. In both species, results did not reveal any significant attraction or behavioral reaction toward DMS when compared to a control chemical stimulus, apart from a short-distance response in bottlenose dolphins. These results suggest that while odontocetes may perceive DMS in water, it apparently does not play a significant role in their foraging ecology. Testing potentially more attractive compounds such as prey extracts with the present method and analyzing surface, underwater and acoustic responses would provide further insights on odontocete feeding behavior. It would also provide valuable clues to studies on the anatomical structures involved in their chemosenses.

Long-finned pilot whales (LFPW) are cetaceans with strong social groups often involved in mass strandings worldwide. However, these beachings occur for reasons that are not fully understood. In 2016, 124 LFPW were stranded on the Chilean Patagonian islands, offering a unique opportunity to obtain crucial information on the ecology, biology, and genetics of this population. In addition, we examined whether persistent organic pollutants (POPs) and trace elements (TEs) were responsible for this mass mortality. Stable isotopes (δ13C & δ15N) and genetic analyses were used to reconstruct the trophic ecology, social structure, and kinship of LFPW and compared to POPs and TEs levels found in LFPW. Mitochondrial DNA analyses on 71 individuals identified four maternal lineages within the stranded LFPW. Of these animals, 32 individuals were analyzed for a suite of POPs, TEs, and lipid content in blubber. The highest levels were found for ΣDDXs (6 isomers) (542.46 ± 433.46 ng/g, lw) and for total Hg (2.79 ± 1.91 mg/kg, dw). However, concentrations found in these LFPW were lower than toxicity thresholds and those reported for LFPW stranded in other regions. Evidence was found of ΣDDX, Σ7PCBs, and Cd bioaccumulation and maternal transfer of POPs in mother/offspring groups. Nevertheless, no clear relationship between contaminant concentrations and LFPW mortality was established. Further research is still needed to assess LFPW populations including conservations status and exposure to chemicals in remote areas such as Patagonia.

Mercury is a neurotoxic chemical that represents one of the greatest pollution threats to Arctic ecosystem health. Evaluating the direct neurotoxic effects of mercury in free ranging wildlife is challenging, necessitating the use of neurochemical biomarkers to assess potential sub-clinical neurological changes. The objective of this study was to characterize the distribution and speciation of mercury, as well as exposure-associated changes in neurochemistry, across multiple brain regions (n = 10) and marine mammal species (n = 5) that each occupy a trophic niche in the Arctic ecosystem. We found consistent species differences in mean brain and brain region-specific concentrations of total mercury (THg) and methyl mercury (MeHg), with higher concentrations in toothed whales (narwhal, pilot whales and harbour porpoise) compared to fur-bearing mammals (polar bear and ringed seal). Mean THg (μg/g dw) in decreasing rank order was: pilot whale (11.9) > narwhal (7.7) > harbour porpoise (3.6) > polar bear (0.6) > ringed seal (0.2). The higher THg concentrations in toothed whales was associated with a marked reduction in the percentage of MeHg (70 %) that had lower brain THg concentrations. This pattern in mercury concentration and speciation corresponded broadly to an overall higher number of mercury-associated neurochemical biomarker correlations in toothed whales. Of the 226 correlations between mercury and neurochemical biomarkers across brain regions, we found 60 (27 %) meaningful relationships (r>0.60 or p < 0.10). We add to the growing weight of evidence that wildlife accumulate mercury in their brains and demonstrate that there is variance in accumulation across species as well as across distinct brain regions, and that some of these exposures may be associated with sub-clinical changes in neurochemistry.

Members of the Delphinidae family are widely distributed across the world's oceans. We used a viral metagenomic approach to identify viruses in orca (Orcinus orca) and short-finned pilot whale (Globicephala macrorhynchus) muscle, kidney, and liver samples from deceased animals. From orca tissue samples (muscle, kidney, and liver), we identified a novel polyomavirus (Polyomaviridae), three cressdnaviruses, and two genomoviruses (Genomoviridae). In the short-finned pilot whale we were able to identify one genomovirus in a kidney sample. The presence of unclassified cressdnavirus within two samples (muscle and kidney) of the same animal supports the possibility these viruses might be widespread within the animal. The orca polyomavirus identified here is the first of its species and is not closely related to the only other dolphin polyomavirus previously discovered. The identification and verification of these viruses expands the current knowledge of viruses that are associated with the Delphinidae family.

Funding: This research was supported by the Office of Naval Research (https://www.onr.navy.mil/) grant N00014-16-1-2858: "PCoD+: Developing widely-applicable models of the population consequences of disturbance” awarded to SMRU Consulting Ltd (http://www.smruconsulting.com/). VH and AMdR benefitted from funding from the European Research Council (https://erc.europa.eu/) under the European Union’s Seventh Framework Programme (FP/2007-2013) / ERC Grant Agreement No. 322814 awarded to AMdR. Nonlethal disturbance of animals can cause behavioral and physiological changes that affect individual health status and vital rates, with potential consequences at the population level. Predicting these population effects remains a major challenge in ecology and conservation. Monitoring fitness-related traits may improve detection of upcoming population changes, but the extent to which individual traits are reliable indicators of disturbance exposure is not well understood, especially for populations regulated by density dependence. Here we study how density dependence affects a population’s response to disturbance and modifies the disturbance effects on individual health and vital rates. We extend an energy budget model for a medium-sized cetacean (the long-finned pilot whale Globicephala melas) to an individual-based population model in which whales feed on a self-replenishing prey base and disturbance leads to cessation of feeding. In this coupled predator-prey system, the whale population is regulated through prey depletion and the onset of yearly repeating disturbances on the whale population at carrying capacity decreased population density and increased prey availability due to reduced top-down control. In populations faced with multiple days of continuous disturbance each year, female whales that were lactating their first calf experienced increased mortality due to depletion of energy stores. However, increased prey availability led to compensatory effects and resulted in a subsequent improvement of mean female body condition, mean age at first reproduction and higher age-specific reproductive output. These results indicate that prey-mediated density dependence can mask negative effects of disturbance on fitness-related traits and vital rates, a result with implications for the monitoring and management of marine mammal populations. Publisher PDF

Summary Cetacean morbillivirus (CeMV) is an important global cause of morbidity and mortality in cetacean populations, with four pathological presentations including non-suppurative encephalitis. We describe an unusual case of dolphin morbillivirus (DMV)-associated non-suppurative encephalitis in a long-finned pilot whale (Globicephala melas), in which the lesions were orientated on the periventricular white matter and comprised prominent multifocal syncytia formation in the absence of systemic lesions. DMV RNA was detected in brain tissue by qRT-PCR and immunohistochemistry for morbillivirus antigen yielded intense labelling of syncytia in periventricular sites, with sparse involvement of the deeper neuroparenchyma. The pattern of lesions raises the possibility of viral dissemination through the cerebrospinal fluid, as described for canine distemper virus, suggesting that similar pathogenic mechanisms may be implicated in lesion development. Further investigation is required to establish the pathogenesis of CeMV encephalitis and the behaviour of the virus within the central nervous system of cetaceans.

Funding: E.P., V.H., J.H., L.N., and D.P.C. were supported by Office of Naval Research (ONR) grant N00014-16-1-2858 (“PCoD+: Developing Widely-Applicable Models of the Population Consequences of Disturbance”). D.P.C. was supported by the Exploration and Production Sound and Marine Life Joint Industry Project of the International Association of Oil and Gas Producers. A.M.d.R. and V.H. were funded by European Research Council Grant no. 322814 awarded to A.M.d.R. M.M. was supported by ONR grant N00014-19-2494. Animals initiate, interrupt, or invest resources in reproduction in light of their physiology and the environment. The energetic risks entailed in an individual’s reproductive strategy can influence the ability to cope with additional stressors, such as anthropogenic climate change and disturbance. To explore the trade-offs between internal state, external resource availability, and reproduction, we applied state-dependent life-history theory (SDLHT) to a dynamic energy budget (DEB) model for long-finned pilot whales (Globicephala melas). We investigated the reproductive strategies emerging from the interplay between fitness maximization and propensity to take energetic risks, as well as the resulting susceptibility of individual vital rates to disturbance. Without disturbance, facultative reproductive behavior from SDLHT and fixed rules in the DEB model led to comparable individual fitness. However, under disturbance, the reproductive strategies emerging from SDLHT increased vulnerability to energetic risks, resulting in lower fitness than fixed rules. These fragile strategies might therefore be unlikely to evolve in the first place. Heterogeneous resource availability favored more cautious (and thus more robust) strategies, particularly when knowledge of resource variation was accurate. Our results demonstrate that the assumptions regarding the dynamic trade-offs underlying an individual’s decision-making can have important consequences for predicting the effects of anthropogenic stressors on wildlife populations. Publisher PDF

This research was funded by Office of Naval Research awards [N00014-08-1-0984, N00014-10-1-0355, N00014-15-1-2341], the Dutch Royal Navy (Koninklijke Nederlandse Marine) and the Norwegian Navy (Forsvarets Forskningsinstitutt – FFI). Group-living animals must communicate to stay in contact. In long-finned pilot whales, there is a trade-off between the benefits of foraging individually at depth and the formation of tight social groups at the surface. Using theoretical modelling and empirical data of tagged pairs within a group, we examined the potential of pilot whale social calls to reach dispersed group members during foraging periods. Both theoretical predictions and empirical data of tag pairs showed a potential for communication between diving and non-diving group members over separation distances up to 385 m (empirical) and 1800 m (theoretical). These distances match or exceed pilot whale dive depths recorded across populations. Call characteristics and environmental characteristics were analysed to investigate determinants of call detectability. Longer calls with a higher sound pressure level (SPL) that were received in a quieter environment were more often detected than their shorter, lower SPL counterparts within a noisier environment. In a noisier environment, calls were louder and had a lower peak frequency, indicating mechanisms for coping with varying conditions. However, the vulnerability of pilot whales to anthropogenic noise is still of concern as the ability to cope with increasing background noise may be limited. Our study shows that combining propagation modelling and actual tag recordings provides new insights into the communicative potential for social calls in orientation and reunion with group members for deep-diving pilot whales. Publisher PDF

The matrilineal long-finned pilot whale presents an antitropical distribution and is divided into two subspecies, one in the temperate seas of the Southern Hemisphere and the other restricted to the North Atlantic and Mediterranean. Until now, population genetic and phylogeographic studies have included localities of most of its Northern Hemisphere distribution, while only the southwestern Pacific has been sampled in the Southern Hemisphere. We add new genetic data from the southeastern Pacific to the published sequences. Low mitochondrial and nuclear diversity was encountered in this new area, as previously reported for other localities. Four haplotypes were found with only one new for the species. Fifteen haplotypes were detected in the global dataset, underlining the species’ low diversity. As previously reported, the subspecies shared two haplotypes and presented a strong phylogeographic structure. The extant distribution of this species has been related to dispersal events during the Last Glacial Maximum. Using the genetic data and Approximate Bayesian Calculations, this study supports this historical biogeographic scenario. From a taxonomic perspective, even if genetic analyses do not support the subspecies category, this study endorses the incipient divergence process between hemispheres, thus maintaining their status and addressing them as Demographically Independent Populations is recommended.

This study presents and evaluates several methods for automated species-level classification of echolocation clicks from three beaked whale species recorded in the northern Gulf of Mexico. The species included are Cuvier's and Gervais' beaked whales, as well as an unknown species denoted Beaked Whale Gulf. An optimal feature set for discriminating the three click types while also separating detected clicks from unidentified delphinids was determined using supervised step-wise discriminant analysis. Linear and quadratic discriminant analyses both achieved error rates below 1% with three features, determined by tenfold cross validation. The waveform fractal dimension was found to be a highly ranked feature among standard spectral and temporal parameters. The top-ranking features were Higuchi's fractal dimension, spectral centroid, Katz's fractal dimension, and -10 dB duration. Six clustering routines, including four popular network-based algorithms, were also evaluated as unsupervised classification methods using the selected feature set. False positive rates of 0.001 and 0.024 were achieved by Chinese Whispers and spectral clustering, respectively, across 200 randomized trials. However, Chinese Whispers clustering yielded larger false negative rates. Spectral clustering was further tested on clicks from encounters of beaked, sperm, and pilot whales in the Tongue of the Ocean, Bahamas.

The US Office of Naval Research and Strategic Environmental Research and Development Program (SERDP) supported the fieldwork as a part of the 3S study collaboration. This study was also supported by the program Bio-Logging Science of the University of Tokyo (UTBLS). To maximize foraging duration at depth, diving mammals are expected to use the lowest cost optimal speed during descent and ascent transit and to minimize the cost of transport by achieving neutral buoyancy. Here, we outfitted 18 deep-diving long-finned pilot whales with multi-sensor data loggers and found indications that their diving strategy is associated with higher costs than those of other deep-diving toothed whales. Theoretical models predict that optimal speed is proportional to (basal metabolic rate/drag)1/3 and therefore to body mass0.05. The transit speed of tagged animals (2.7±0.3 m s−1) was substantially higher than the optimal speed predicted from body mass (1.4–1.7 m s−1). According to the theoretical models, this choice of high transit speed, given a similar drag coefficient (median, 0.0035) to that in other cetaceans, indicated greater basal metabolic costs during diving than for other cetaceans. This could explain the comparatively short duration (8.9±1.5 min) of their deep dives (maximum depth, 444±85 m). Hydrodynamic gliding models indicated negative buoyancy of tissue body density (1038.8± 1.6 kg m–3, ±95% credible interval, CI) and similar diving gas volume (34.6±0.6 ml kg−1, ±95% CI) to those in other deep-diving toothed whales. High diving metabolic rate and costly negative buoyancy imply a ‘spend more, gain more’ strategy of long-finned pilot whales, differing from that in other deep-diving toothed whales, which limits the costs of locomotion during foraging. We also found that net buoyancy affected the optimal speed: high transit speeds gradually decreased during ascent as the whales approached neutral buoyancy owing to gas expansion. Publisher PDF

The knowledge of the vocal repertoire of pilot whales is very limited. In this paper, the vocal repertoire of long-finned pilot whales recorded during different encounters in the Vestfjord in northern Norway between November 2006 and August 2010 are described. Sounds were analysed using two different methods: (1) an observer-based audio-visual inspection of FFT-derived spectrograms, with which, besides a general variety of clicks, buzzes, nonharmonic sounds, and whistles, 129 different distinct call types and 25 subtypes were distinguished. These call types included pulsed calls and discrete structured whistles varying from simple to highly complex structures composed of several segments and elements. In addition, ultrasonic whistles previously not described for pilot whales were found. In addition to the diversity of single calls, call sequences consisting of repetitions and combinations of specific call types were recorded and (2) a parametric approach that permitted the confirmation of the high variability in pilot whale call structures was developed. It is concluded that the pilot whale vocal repertoire is among the most complex for the mammalian species and the high structural variability, along with call repetitions and combinations, require a closer investigation to judge their importance for vocal communication.

Echolocating toothed whales produce powerful clicks pneumatically to detect prey in the deep sea where this long-range sensory channel makes them formidable top predators. However, air supplies for sound production compress with depth following Boyle’s law suggesting that deep-diving whales must use very small air volumes per echolocation click to facilitate continuous sensory flow in foraging dives. Here we test this hypothesis by analysing click-induced acoustic resonances in the nasal air sacs, recorded by biologging tags. Using 27000 clicks from 102 dives of 23 tagged pilot whales (Globicephala macrorhynchus), we show that click production requires only 50 μL of air/click at 500 m depth increasing gradually to 100 μL at 1000 m. With such small air volumes, the metabolic cost of sound production is on the order of 40 J per dive which is a negligible fraction of the field metabolic rate. Nonetheless, whales must make frequent pauses in echolocation to recycle air between nasal sacs. Thus, frugal use of air and periodic recycling of very limited air volumes enable pilot whales, and likely other toothed whales, to echolocate cheaply and almost continuously throughout foraging dives, providing them with a strong sensory advantage in diverse aquatic habitats. Echolocating toothed whales produce powerful clicks pneumatically to detect prey in the deep sea where this long-range sensory channel makes them formidable top predators. However, air supplies for sound production compress with depth following Boyle’s law suggesting that deep-diving whales must use very small air volumes per echolocation click to facilitate continuous sensory flow in foraging dives. Here we test this hypothesis by analysing click-induced acoustic resonances in the nasal air sacs, recorded by biologging tags. Using 27000 clicks from 102 dives of 23 tagged pilot whales (Globicephala macrorhynchus), we show that click production requires only 50 µL of air/click at 500 m depth increasing gradually to 100 µL at 1000 m. With such small air volumes, the metabolic cost of sound production is on the order of 40 J per dive which is a negligible fraction of the field metabolic rate. Nonetheless, whales must make frequent pauses in echolocation to recycle air between nasal sacs. Thus, frugal use of air and periodic recycling of very limited air volumes enable pilot whales, and likely other toothed whales, to echolocate cheaply and almost continuously throughout foraging dives, providing them with a strong sensory advantage in diverse aquatic habitats.

High exposures of mammalian species to inorganic mercury (HgII) and methylmercury (MeHg) have been associated with adverse effects on behavior and reproduction. Different mammalian species exhibit varying responses to similar external exposure levels, reflecting potential differences in Hg toxicokinetics. Here, we use Hg stable isotopes, total Hg, MeHg and selenium (Se) concentrations measured in multiple tissues of North Atlantic pilot whales (Globicephala melas) to investigate processes affecting the distribution and accumulation of HgII and MeHg. We find that simple mixing of two distinct isotopic end-members: MeHg (1.4‰) and HgII (-1.6‰) can explain the observed variability of δ202Hg in brain tissue. A similar isotopic composition for the MeHg end-member in the brain, muscle, heart, and kidney suggests efficient exchange of MeHg in blood throughout the body. By contrast, the Hg isotopic composition of the liver of adult whales is different from younger whales and other tissues that follow the two-end member mixing model. Measured Se:Hg ratios are lowest in adult whales with the highest levels of MeHg exposure. In these individuals, Se availability is likely reduced by complexation with demethylated HgII. We speculate that this results in a higher fraction of labile HgII eliminated from the liver of adult whales compared to young whales and subsequent redistribution to other tissues, potentially affecting toxicity.

Research funding was provided by the US Office of Naval Research, the DGA/TN (France), the UK Natural Environmental Research Council, and the Ministries of Defence of Norway and The Netherlands. PLT acknowledges funding received from the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland). MASTS is funded by the Scottish Funding Council (Grant reference HR09011) and contributing institutions. CC acknowledges statistical support provided by the Multi-study OCean acoustics Human effects Analysis (MOCHA) project funded by the United States Office of Naval Research (Grant N00014-12-1-0204). Killer whales (KW) may be predators or competitors of other cetaceans. Since their foraging behavior and acoustics differ among populations ('ecotypes'), we hypothesized that other cetaceans can eavesdrop on KW sounds and adjust their behavior according to the KW ecotype. We performed playback experiments on long-finned pilot whales (Globicephala melas) in Norway using familiar fish-eating KW sounds (fKW) simulating a sympatric population that might compete for foraging areas, unfamiliar mammal-eating KW sounds (mKW) simulating a potential predator threat, and two control sounds. We assessed behavioral responses using animal-borne multi-sensor tags and surface visual observations. Pilot whales barely changed behavior to a broadband noise (CTRL-), whereas they were attracted and exhibited spyhops to fKW, mKW, and to a repeated-tonal upsweep signal (CTRL+). Whales never stopped nor started feeding in response to fKW, whereas they reduced or stopped foraging to mKW and CTRL+. Moreover, pilot whales joined other subgroups in response to fKW and CTRL+, whereas they tightened individual spacing within group and reduced time at surface in response to mKW. Typical active intimidation behavior displayed to fKW might be an antipredator strategy to a known low-risk ecotype or alternatively a way of securing the habitat exploited by a heterospecific sympatric population. Cessation of feeding and more cohesive approach to mKW playbacks might reflect an antipredator behavior towards an unknown KW ecotype of potentially higher risk. We conclude that pilot whales are able to acoustically discriminate between familiar and unfamiliar KW ecotypes, enabling them to adjust their behavior according to the perceived disturbance type. Postprint

This research was supported by the Office of Naval Research grant N00014-16-1-2858: "PCoD+: Developing widely-applicable models of the population consequences of disturbance”. VH and AMdR benefitted from funding from the European Research Council under the European Union’s Seventh Framework rogramme (F /2007-2013) / ERC Grant Agreement No. 322814 awarded to AMdR. Understanding the full scope of human impact on wildlife populations requires a framework to assess the population‐level repercussions of nonlethal disturbance. The Population Consequences of Disturbance (PCoD) framework provides such an approach, by linking the effects of disturbance on the behavior and physiology of individuals to their population‐level consequences. Bio‐energetic models have been used as implementations of PCoD, as these integrate the behavioral and physiological state of an individual with the state of the environment, to mediate between disturbance and biological significant changes in vital rates (survival, growth, and reproduction). To assess which levels of disturbance lead to adverse effects on population growth rate requires a bio‐energetic model that covers the complete life cycle of the organism under study. In a density‐independent setting, the expected lifetime reproductive output of a single female can then be used to predict the level of disturbance that leads to population decline. Here, we present such a model for a medium‐sized cetacean, the long‐finned pilot whale (Globicephala melas). Disturbance is modeled as a yearly recurrent period of no resource feeding for the pilot whale female and her calf. Short periods of disturbance lead to the pre‐weaned death of the first one or more calves of the young female. Higher disturbance levels also affect survival of calves produced later in the life of the female, in addition to degrading female survival. The level of disturbance that leads to a negative population growth rate strongly depends on the available resources in the environment. This has important repercussion for the timing of disturbance if resource availability fluctuates seasonally. The model predicts that pilot whales can tolerate on average three times longer periods of disturbance in seasons of high resource availability, compared to disturbance happening when resources are low. Although our model is specifically parameterized for pilot whales, it provides useful insights into the general consequences of nonlethal disturbance. If appropriate data on life history and energetics are available, it can be used to provide management advice for specific species or populations. Publisher PDF

Whales accumulate mercury (Hg), but do not seem to show immediate evidence of toxic effects. Analysis of different tissues (liver, kidney, muscle) and biofluids (blood, milk) from a pod of stranded long-finned pilot whales (Globicephala melas) showed accumulation of Hg as a function of age, with a significant decrease in the MeHg fraction. Isotopic analysis revealed remarkable differences between juvenile and adult whales. During the first period of life, Hg in the liver became isotopically lighter (δ202Hg decreased) with a strongly decreasing methylmercury (MeHg) fraction. We suggest this is due to preferential demethylation of MeHg with the lighter Hg isotopes and transport of MeHg to less sensitive organs, such as the muscles. Also changes in diet, with high MeHg intake in utero and during lactation, followed by increasing consumption of solid food contribute to this behavior. Interestingly, this trend in δ202Hg is reversed for livers of adult whales (increasing δ202Hg value), accompanied by a progressive decrease of δ202Hg in muscle at older ages. These total Hg (THg) isotopic trends suggest changes in the Hg metabolism of the long-finned pilot whales, development of (a) detoxification mechanism(s) (e.g., though the formation of HgSe particles), and Hg redistribution across the different organs.

International audience; New data are reported from analyses of stomach contents from 114 long-finned pilot whalesmass-stranded at four locations around Tasmania, Australia from 1992–2006. Identifiableprey remains were recovered from 84 (74%) individuals, with 30 (26%) individuals (17females and 13 males) having empty stomachs. Prey remains comprised 966 identifiablelower beaks and 1244 upper beaks, belonging to 17 families (26 species) of cephalopods.Ommastrephidae spp. were the most important cephalopod prey accounting for 16.9% bynumber and 45.6% by reconstructed mass. Lycoteuthis lorigera was the next most important,followed by Ancistrocheirus lesueurii. Multivariate statistics identified significant differencesin diet among the four stranding locations. Long-finned pilot whales foraging offSouthern Australia appear to be targeting a diverse assemblage of prey (�10 species dominatedby cephalopods). This is compared to other similar studies from New Zealand andsome locations in the Northern Hemisphere, where the diet has been reported to be primarilyrestricted to �3 species dominated by cephalopods. This study emphasises the importanceof cephalopods as primary prey for Southern long-finned pilot whales and othermarine vertebrates, and has increased our understanding of long-finned pilot whale diet inSouthern Ocean waters.

Background Adequate thyroid function during pregnancy is essential for optimal fetal growth. Gestational exposure to perfluoroalkyl substances (PFAS) can negatively affect birth size and disrupt maternal and neonatal thyroid function, although the interrelationship is unclear. Objective We aimed to quantify the associations between maternal serum–PFAS concentrations and birth weight, birth length, and cranial circumference. We also aimed to estimate associations between PFAS and thyroid hormone (TH) concentrations, thereby elucidating whether THs potentially mediate the associations between PFAS concentrations and birth size. Methods We studied a population-based prospective cohort of 172 mother-singleton pairs from the Faroe Islands. Twelve PFAS were measured in maternal serum obtained at 34 weeks of gestation. THs were measured in maternal and cord serum. Associations between PFAS concentrations and birth size and TH concentrations were estimated using multivariable linear regressions. Sex-stratified analyses along with a mediation analysis were performed to estimate potential mediating effects of THs in the association between PFAS and birth outcomes. Results Several PFASs were negatively associated with birth weight, length, and head circumference, and a general positive association between maternal serum–PFASs and cord serum–thyroid-stimulating hormone (TSH; also known as thyrotropin) was found. For instance, a doubling in perfluorooctane sulfonate (PFOS) and perfluorooctanoic acid (PFOA) was associated with a 53% (95% CI, 18%-99%) and 40% (95% CI, 8%-81%) increases in TSH concentrations, respectively. There was little evidence of sexually dimorphic associations. Overall, THs were not found to mediate associations between PFASs and birth size. Conclusion In this study, several PFASs were negatively associated with birth size and increased THs; however, this did not explain lower birth weight among children exposed to PFAS.

The Southern Hemisphere long-finned pilot whale (Globicephala melas edwardii) is a top predator in the New Zealand pelagic food web, feeding predominantly on arrow squids. This study quantified trace element concentrations (Al, As, Cd, Co, Cr, Cu, Fe, Hg, Ni, Mn, Pb, Se, V, Zn) in four tissues (blubber, kidney, liver, muscle) from 21 individuals from stranding sites in New Zealand. Maximum Cd and Hg concentrations were measured in liver and kidney, respectively. Selenium had a positive correlation with Cd and Hg, suggesting the involvement of Se in Cd and Hg detoxification. Arrow squids from the whales' stomach contents were DNA barcoded and identified as Nototodarus sloanii. Trace element concentrations were measured in squid samples from the whale stomach contents. The significant correlation for Hg between the squid tissue and the whale tissue suggests that arrow squids play a major role in trace element uptake by G. m. edwardii.

The bioaccumulation of metals was investigated by analysis of liver, kidney, muscle and brain tissue of a pod of 21 long-finned pilot whales (Globicephala melas) of all ages stranded in Scotland, UK. The results are the first to report cadmium (Cd) passage through the blood-brain barrier of pilot whales and provide a comprehensive study of the long-term (up to 35 years) mammalian exposure to the environmental pollutants. Additionally, linear accumulation of mercury (Hg) was observed in all studied tissues, whereas for Cd this was only observed in the liver. Total Hg concentration above the upper neurochemical threshold was found in the sub-adult and adult brains and methylmercury (MeHg) of 2.2mg/kg was found in the brain of one individual. Inter-elemental analysis showed significant positive correlations of Hg with selenium (Se) and Cd with Se in all studied tissues. Furthermore, differences in the elemental concentrations in the liver and brain tissues were found between juvenile, sub-adult and adult groups. The highest concentrations of manganese, iron, zinc, Se, Hg and MeHg were noted in the livers, whereas Cd predominantly accumulated in the kidneys. High concentrations of Hg and Cd in the tissues of pilot whales presented in this study reflect ever increasing toxic stress on marine mammals.

The use of trace metals and rare-earth elements (REEs) is increasing in the mining, metallurgic, electronic, and automobile industries due to their magnetic, heat-resistant, and phosphorescent properties. While large amounts of these metals are released to the environment, the toxic consequences in marine organisms are poorly understood. In Chile a mass stranding event of long-finned pilot whales (LFPW) (Globicephala melas) occurred in 2016 due to unknown consequences. Al, Ce, Cr, Cu, Tl, and Zn concentrations were analyzed in LFPW blubber tissue and correlated with body size and age class of individuals. While Al and Zn were higher in juvenile individuals, Ce, Cu, Cr, and Tl were higher in adults. This study provides the first base line of trace metals and REE in LFPW from the southern hemisphere and demonstrates the existence and persistence of trace elements in marine top predators from remote ecosystems like the Chilean Patagonia.

Funding:Commander, U.S. Pacific Fleet Environmental Readiness Division and NMFS Pacific Islands Fisheries Science Center; NMFS West Coast Region; Scripps Institution of Oceanography Edna Bailey Sussman Research Fellowship; and Woods Hole Oceanographic Institution. 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. Postprint

This research was supported by award RC-2154 from the Strategic Environmental Research and Development Program and funding from the Naval Facilities Engineering Command Atlantic and National Oceanic and Atmospheric Administration Fisheries, Southeast Region. The risk of predation is often invoked as an important factor influencing the evolution of social organization in cetaceans, but little direct information is available about how these aquatic mammals respond to predators or other perceived threats. We used controlled playback experiments to examine the behavioral responses of short-finned pilot whales (Globicephala macrorhynchus) off Cape Hatteras, NC, USA, and Risso's dolphins (Grampus griseus) off the coast of Southern California, USA, to the calls of a potential predator, mammal-eating killer whales. We transmitted calls of mammal-eating killer whales, conspecifics and baleen whales to 10 pilot whales and four Risso's dolphins equipped with multi-sensor archival acoustic recording tags (DTAGs). Only playbacks of killer whale calls resulted in significant changes in tagged animal heading. The strong responses observed in both species occurred only following exposure to a subset of killer whale calls, all of which contained multiple non-linear properties. This finding suggests that these structural features of killer whale calls convey information about predatory risk to pilot whales and Risso's dolphins. The observed responses differed between the two species; pilot whales approached the sound source while Risso's dolphins fled following playbacks. These divergent responses likely reflect differences in anti-predator response mediated by the social structure of the two species. Publisher PDF

Faroe Island pilot whales have been documented to have high body burdens of organohalogen contaminants (OHCs), including polychlorinated biphenyls (PCBs) and polybrominated diphenyl ethers (PBDEs), but low burdens of their respective hydroxylated metabolites (OH-PCBs and OH-PBDEs). The present study investigated the hepatic expression and/or catalytic activities of phase I and II biotransformation enzymes in relation to hepatic concentrations of target OHCs, including OH-PCBs and OH-PBDEs, in long-finned pilot whales (Globicephala melas) from the Northeastern Atlantic. CYP1A, 2B, 2E and 3A protein expressions were identified in juveniles and adult males, but not in adult females. Ethoxyresorufin-O-deethylase (EROD) activity was significantly lower in adult females than in juveniles and adult males. Using multivariate analyses to investigate relationships between biological responses and OHC concentrations, a positive relationship was identified between EROD and OHCs. The activity levels of phase II conjugating enzymes (uridine 5′-diphospho-glucuronosyltransferase [UDPGT], and glutathione S-transferase [GST]) were low. The analyses of mRNA Expression did not show correlative relationships with OHC concentrations, but cyp1a and ahr transcripts were positively correlated with EROD activity. We suggest that the low concentrations of OH-PCBs and OH-PBDEs reported in pilot whales is probably due to the identified low phase I biotransformation activities in the species. © 2017. This is the authors’ accepted and refereed manuscript to the article. Locked until 19.12.2019 due to copyright restrictions. This manuscript version is made available under the CC-BY-NC-ND 4.0 license http://creativecommons.org/licenses/by-nc-nd/4.0/