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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.

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.

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.

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.

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.

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.

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/

Fatal Brucella ceti infection with histological lesions specific to the central nervous system has been described in only 3 species of cetaceans: striped dolphins Stenella coeruleoalba, Atlantic white-sided dolphins Lagenorhynchus acutus and short-beaked common dolphins Delphinus delphis. This paper describes the first report of a B. ceti-associated meningoencephalitis in a long-finned pilot whale Globicephala melas, showing the increasing range of species susceptibility. Brucella was recovered in larger numbers from cerebrospinal fluid than from brain tissue and is the sample of choice for isolation.

To the Editor: Four strains of cetacean morbillivirus (CeMV; family Paramyxoviridae, genus Morbillivirus) have been detected in the global cetacean population: porpoise morbillivirus (1), dolphin morbillivirus (2), pilot whale morbillivirus (PWMV) (3), and Longman’s beaked whale morbillivirus (4). In addition, 2 novel CeMV sequences or strains isolated from the Indo-Pacific bottlenose dolphin (Tursiops aduncus) and the Guiana dolphin (Sotalia guianensis) have been recently reported in the Southern Hemisphere (5,6). Pilot whales are known to be susceptible to 2 strains of CeMV, PWMV, and dolphin morbillivirus (3,7,8). Only 2 deaths of whales have been reported to be caused by PWMV: 1 long-finned pilot whale (Globicephala melas) (3) and 1 short-finned pilot whale (G. macrorhynchus) (8). We report deaths of 3 short-finned pilot whales caused by PWMV in the northeastern Atlantic Ocean along the coast of the Canary Islands, Spain. During mid-January–May 2015, a total of 3 whales (animals 1, 2, and 3) were found dead along the coasts of the Canary Islands (Table). Complete standardized necropsy was performed for all whales. Tissue samples from animals 1 and 2 were fixed in 10% neutral-buffered formalin for histologic and immunohistochemical analyses (Technical Appendix Figure). Immunohistochemical analysis was performed (brain, intestine, lymph nodes, lung, kidney, adrenal gland, uterus, ovary, testis, and spleen) by using a monoclonal antibody against the nucleoprotein of canine distemper virus (CDV-NP; VMRD, Inc., Pullman, WA, USA) (7). Samples of lung, pulmonary lymph nodes, larynx, laryngeal tonsil, intestine, spleen, and brain were frozen (−80°C) for virologic analysis. Table Characteristics for 3 short-finned pilot whales stranded along the Canary Islands, Spain, 2015* Grossly, the most remarkable findings in animal 1 were severe suppurative rhinitis, with clogged nasal passages by the accumulation of large quantity of purulent material, otitis media, sacculitis, and laryngitis. Severe diffuse epithelial hyperplasia and hyperkeratosis was observed along the upper respiratory tract and keratinized stomach. Animal 2 had severe proliferative dermatitis and cheilitis, and severe, suppurative, laryngeal tonsillitis. Animal 3 had advanced autolysis, which precluded pathologic analysis. Histologically, moderate, multifocal, bronchointerstitial pneumonia, severe suppurative tonsillitis and systemic lymphoid depletion were identified in animals 1 and 2. Severe nonsuppurative meningoencephalitis with neuronal and glial cell degeneration and necrosis, microgliosis and syncytial cells were observed in animal 2. Immunohistochemical analysis showed morbillivirus antigen in the bronchiolar epithelium, type 2 pneumocytes, and alveolar multinucleate cells. Syncytia from lymph nodes, laryngeal tonsil, spleen, and intestine also showed positive immunolabeling for morbillivirus. Epithelial tropism caused by the virus was suggested by identification of viral antigen in epithelia of the lung, larynx, keratinized stomach, intestine, kidney, urinary bladder, epididymis, and endometrial glands. In addition, intense immunolabeling was detected in neurons (soma, dendrites, axon hillock, and axons) and glial cells, primarily throughout the cerebral gray matter of animal 2. Molecular detection of CeMV was performed by a using a 1-step reverse transcription PCR for a 426-bp conserved region of the phosphoprotein gene (7). All tested samples from animals 1 and 2 and a laryngeal tonsil sample from animal 3 showed positive PCR results. Because co-infections with herpesvirus and morbillivirus were observed during morbillivirus epizootics in seals in 1988 and dolphins in 2006–2007, we also tested the same tissue for herpesvirus by conventional nested PCR (9). Herpesvirus DNA was detected in all samples from animal 1 except lung, although no specific lesions compatible with this infectious agent were observed. A pool containing all morbillivirus-positive PCR amplicons for animals 1 and 2 (GenBank accession nos. {"type":"entrez-nucleotide","attrs":{"text":"KT006289","term_id":"961377517"}}KT006289 and {"type":"entrez-nucleotide","attrs":{"text":"KT006290","term_id":"961377519"}}KT006290), a PCR amplicon for the brain sample from animal 2 (GenBank accession no. {"type":"entrez-nucleotide","attrs":{"text":"KT006291","term_id":"961377521"}}KT006291), and a PCR amplicon for the larynx from animal 3 were sequenced. A BLAST search (http://www.ncbi.nlm.nih.gov/blast/Blast.cgi) showed that amplified samples were nearly identical to reference PWMV sequences (GenBank accession nos. {"type":"entrez-nucleotide","attrs":{"text":"AF200817","term_id":"7329127"}}AF200817 [3] and {"type":"entrez-nucleotide","attrs":{"text":"FJ842381","term_id":"239775410"}}FJ842381 [8]). The sequence obtained from animal 3 was too short and degenerated to be accurately classified as CeMV, although it showed high homology with PWMV and porpoise morbillivirus. It has been proposed that pilot whales might be enzootically infected with CeMV (10). These whales might be responsible for maintaining and transmitting CeMV over long distances or to other odontocetes. No die-offs have been observed in these species. However, an outbreak of a lethal morbillivirus infection in long-finned pilot whales caused by a dolphin morbillivirus strain occurred in the Mediterranean Sea during the end of October 2006–April 2007 (7). Results of this study support the previous hypothesis that pilot whales have a species-adapted morbillivirus but indicate that lethal infections are not as rare as previously believed (3). The tropism of the virus in these cases, the high number of multinucleated syncytial cells, and the severity of the lesions resemble the acute systemic symptoms observed in dolphins infected with morbillivirus (2). Thus, pilot whales in the northeastern Atlantic Ocean could be at risk for infection, especially in one of the main pilot whale–watching regions between La Gomera and Southern Tenerife Islands in the Canary Islands, which has >700,000 visitors each year. Technical Appendix. Microscopic images of tissue samples from 2 short-finned pilot whales (Globicephala macrorhynchus) from the eastern Atlantic Ocean stranded along the Canary Islands, Spain, 2015. Click here to view.(380K, pdf)

Blubber from Faroese pilot whales (Globicephala melas) was analysed for brominated dioxins PBDD/Fs, with a subset also analysed for chlorinated dioxins, PCDD/Fs. The studied individuals were restricted to juvenile male whales sampled in the Faroe Islands during the period 1997-2013. Among the PBDD/Fs, the furans were predominant, although the relative abundance of various congeners differed between samples. Furans accounted for, on average, 79% of the ∑PBDD/Fs in the samples, with 1,2,3,4,6,7,8-HpBDF the most abundant congener, found in half of the analysed pilot whales. The concentration range for ∑PBDD/Fs among the samples was 0.080-71 pg/g l.w. (lipid weight), and the sum of toxic equivalents ranged from 0.0039 to 4.7 pg TEQ/g l.w. No relationship was found between PBDD/Fs and PCDD/Fs. In addition, 20 pilot whale samples from the period 2010-2013 were analysed for PBDEs. Several PBDE congeners were found in all of the sampled pilot whales, and at noticeably higher levels than PBDD/Fs and PCDD/Fs. The ∑PBDEs ranged from 140 to 1900 ng/g l.w., with BDE #47 the most abundant congener detected in the samples. Results from the present study were then compared with data from previous studies on pilot wales to investigate temporal trends between 1986 and 2013. The comparison indicated that PBDE concentrations in juvenile males have decreased from 1996 to the latest observations in 2013. No relationship between the concentration levels of PBDD/Fs and PBDEs in the sampled pilot whales could be identified, which indicates possible differences in the metabolism of, or exposure to, PBDEs and PBDD/Fs.

Poly- and perfluoroalkyl substances (PFASs) are persistent, bioaccumulative anthropogenic compounds associated with adverse health impacts on humans and wildlife. PFAS production changed in North America and Europe around the year 2000, but impacts on wildlife appear to vary across species and location. Unlike other mammal species, cetaceans lack the enzyme for transforming an important intermediate precursor (perfluorooctane sulfonamide: FOSA), into a prevalent compound in most wildlife (perfluorooctane sulfonate: PFOS). Thus, their tissue burden differentiates these two compounds while other mammals contain PFOS from both direct exposure and precursor degradation. Here we report temporal trends in 15 PFASs measured in muscle from juvenile male North Atlantic pilot whales (Globicephala melas) harvested between 1986 and 2013. FOSA accounted for a peak of 84% of the 15 PFASs around 2000 but declined to 34% in recent years. PFOS and long-chained PFCAs (C9-C13) increased significantly over the whole period (2.8% yr(−1) to 8.3% yr(−1)), but FOSA declined by 13% yr(−1) after 2006. Results from FOSA partitioning and bioaccumulation modeling forced by changes in atmospheric inputs reasonably capture magnitudes and temporal patterns in FOSA concentrations measured in pilot whales. Rapid changes in atmospheric FOSA in polar and subpolar regions around 2000 helps to explain large declines in PFOS exposure for species that metabolize FOSA, including seafood consuming human populations. This work reinforces the importance of accounting for biological exposures to PFAS precursors.

Mass strandings of whales and dolphins have puzzled biologists since Aristotle. Although environmental factors are often assumed to initiate strandings, social forces must also influence the dynamics of many of these events, particularly for the primary species involved in mass strandings, the long-finned pilot whales ( Globicephala melas). Here, we test two hypotheses derived from common assumptions about the social dynamics of long-finned pilot whales by identifying maternal lineages from mtDNA haplotypes and inferring kinship from microsatellite genotypes of 490 individuals from 12 stranding events. Contrary to the “extended matriline” hypothesis, we found that multiple maternal lineages were present in at least 9 of the 12 mass strandings. Contrary to the “kinship cohesion” hypothesis, we found no correlation between spatial distribution and kinship along the stranding beach. Most notably, we documented the spatial disruption of the expected proximity between mothers and their dependent calves. These results challenge the common assumption that kinship-based behavior, such as care-giving, are a primary factor in these mass strandings. We suggest instead that disruption of kinship bonds could result from interactions among unrelated social groups during feeding or mating aggregations, perhaps playing a causal role in these events. Our finding that dependent calves were often spatially separated or absent from their mothers has important implications for humane management of rescue efforts. To improve our understanding of the social causes and consequences of mass strandings, future documentation of strandings should include exhaustive DNA sampling, with accompanying spatial and temporal records.

Pilot whales are two cetacean species (Globicephala melas and G. macrorhynchus) whose distributions are correlated with water temperature and partially overlap in some areas like the North Atlantic Ocean. In the context of global warming, distribution range shifts are expected to occur in species affected by temperature. Consequently, a northward displacement of the tropical pilot whale G. macrorynchus is expected, eventually leading to increased secondary contact areas and opportunities for interspecific hybridization. Here, we describe genetic evidences of recurrent hybridization between pilot whales in northeast Atlantic Ocean. Based on mitochondrial DNA sequences and microsatellite loci, asymmetric introgression of G. macrorhynchus genes into G. melas was observed. For the latter species, a significant correlation was found between historical population growth rate estimates and paleotemperature oscillations. Introgressive hybridization, current temperature increases and lower genetic variation in G. melas suggest that this species could be at risk in its northern range. Under increasing environmental and human-mediated stressors in the North Atlantic Ocean, it seems recommendable to develop a conservation program for G. melas.

Cetacean karyotypes possess exceptionally stable diploid numbers and highly conserved chromosomes. To date, only toothed whales (Odontoceti) have been analyzed by comparative chromosome painting. Here, we studied the karyotype of a representative of baleen whales, the gray whale (Eschrichtius robustus, Mysticeti), by Zoo-FISH with dromedary camel and human chromosome-specific probes. We confirmed a high degree of karyotype conservation and found an identical order of syntenic segments in both branches of cetaceans. Yet, whale chromosomes harbor variable heterochromatic regions constituting up to a third of the genome due to the presence of several types of repeats. To investigate the cause of this variability, several classes of repeated DNA sequences were mapped onto chromosomes of whale species from both Mysticeti and Odontoceti. We uncovered extensive intrapopulation variability in the size of heterochromatic blocks present in homologous chromosomes among 3 individuals of the gray whale by 2-step differential chromosome staining. We show that some of the heteromorphisms observed in the gray whale karyotype are due to distinct amplification of a complex of common cetacean repeat and heavy satellite repeat on homologous autosomes. Furthermore, we demonstrate localization of the telomeric repeat in the heterochromatin of both gray and pilot whale (Globicephala melas, Odontoceti). Heterochromatic blocks in the pilot whale represent a composite of telomeric and common repeats, while heavy satellite repeat is lacking in the toothed whale consistent with previous studies.

A selection of MeO-BDE and BDE congeners were analyzed in pooled blubber samples of pilot whale (Globicephala melas), ringed seal (Phoca hispida), minke whale (Balaenoptera acutorostrata), fin whale (Balaenoptera physalus), harbor porpoise (Phocoena phocoena), hooded seal (Cystophora cristata), and Atlantic white-sided dolphin (Lagenorhynchus acutus), covering a time period of more than 20 years (1986-2009). The analytes were extracted and cleaned-up using open column extraction and multi-layer silica gel column chromatography. The analysis was performed using both low resolution and high resolution GC-MS. MeO-PBDE concentrations relative to total PBDE concentrations varied greatly between sampling periods and species. The highest MeO-PBDE levels were found in the toothed whale species pilot whale and white-sided dolphin, often exceeding the concentration of the most abundant PBDE, BDE-47. The lowest MeO-PBDE levels were found in fin whales and ringed seals. The main MeO-BDE congeners were 6-MeO-BDE47 and 2'-MeO-BDE68. A weak correlation only between BDE47 and its methoxylated analog 6-MeO-BDE47 was found and is indicative of a natural source for MeO-PBDEs.

SUMMARY Long-finned pilot whales are highly social odontocetes found in temperate and subpolar regions. This species is particularly known for its interaction with fisheries as well as its mass strandings. Recent tagging work has provided some information about pilot whales in the wild but, even though they have been successfully kept in captivity, little is known about their sensory capabilities. This study investigates the hearing abilities of a rehabilitated 2 year old male long-finned pilot whale. A complete audiogram was collected using auditory evoked potential techniques that included measurements of nine frequencies from 4 to 100 kHz presented as sinusoidally amplitude-modulated tones. The results indicated that the region of best hearing was between 11.2 and 50 kHz and the subject had relatively poor high frequency hearing compared with other odontocete species. This study emphasizes the importance of collecting basic hearing measurements from new species, understanding diagnostic life histories as well as continuously increasing the sample size of audiometry measurements within and between odontocete species as animals become available.

Recordings of animal vocalization can lack information about sender and context. This is often the case in studies on marine mammals or in the increasing number of automated bioacoustics monitorings. Here, we develop a framework to estimate group specificity without specific sender information. We introduce and apply a bag-of-calls-and-coefficients approach (BOCCA) to study ensembles of cepstral coefficients calculated from vocalization signals recorded from a given animal group. Comparing distributions of such ensembles of coefficients by computing relative entropies reveals group specific differences. Applying the BOCCA to ensembles of calls recorded from group of long-finned pilot whales in northern Norway, we find that differences of vocalizations within social groups of pilot whales (Globicephala melas) are significantly lower than intergroup differences.

The pilot whale Globicephala melas, the sperm whale Physeter macrocephalus, and the fin whale Balaenoptera physalus are large cetaceans permanently inhabiting the Mediterranean Sea. These species are subjected to numerous anthropogenic threats such as exposure to high levels of contaminants. Therefore, selected persistent organic pollutants POPs (29 PCBs, 15 organochlorine compounds, 9 PBDEs and 17 PCDD/Fs) were analysed in blubber biopsies of 49 long-finned pilot whales, 61 sperm whales and 70 fin whales sampled in the North Western Mediterranean Sea (NWMS) from 2006 to 2013. Contamination profile and species feeding ecology were then combined through the use of stable isotopes. δ(13)C, δ(15)N values and POPs levels were assessed through IR-MS and GC-MS respectively. To assess the toxic potency of the dioxin-like compounds, the TEQ approach was applied. δ(15)N values were 12.2±1.3‰ for sperm whales, 10.5±0.7‰ for pilot whales and 7.7±0.8‰ in fin whales, positioning sperm whales at higher trophic levels. δ(13)C of the two odontocetes was similar and amounted to -17.3±0.4‰ for sperm whales and -17.8±0.3‰ for pilot whales; whilst fin whales were more depleted (-18.7±0.4‰). This indicates a partial overlap in toothed-whales feeding habitats, while confirms the differences in feeding behaviour of the mysticete. Pilot whales presented higher concentrations than sperm whales for ΣPCBs (38,666±25,731 ng g(-1)lw and 22,849±15,566 ng g(-1) lw respectively), ΣPBDEs (712±412 ng g(-1) lw and 347±173 ng g(-1) lw respectively) and ΣDDTs (46,081±37,506 ng g(-1) lw and 37,647±38,518 ng g(-1) lw respectively). Fin whales presented the lowest values, in accordance with its trophic position (ΣPCBs: 5721±5180 ng g(-1) lw, ΣPBDEs: 177±208 ng g(-1) lw and ΣDDTs: 6643±5549 ng g(-1) lw). Each species was characterized by large inter-individual variations that are more related to sex than trophic level, with males presenting higher contaminant burden than females. The discriminant analysis (DA) confirmed how DDTs and highly chlorinated PCBs were influential in differentiating the three species. Pollutant concentrations of our species were significantly higher than both their Southern Hemisphere and North Atlantic counterparts, possibly due to the particular Mediterranean geomorphology, which influences pollutants distribution and recycle. Dioxin-like PCBs accounted for over 80% of the total TEQ. This study demonstrated (1) an important exposure to pollutants of Mediterranean cetaceans, often surpassing the estimated threshold toxicity value of 17,000 ng g(-1) lw for blubber in marine mammals; and (2) how the final pollutant burden in these animals is strongly influenced not only by the trophic position but also by numerous other factors such as sex, age, body size and geographical distribution.

Concentrations of PCBs, organochlorine pesticides (OCPs), brominated flame retardants and a suite of relevant metabolites of these POPs, in all 175 different compounds, were determined in liver and plasma of traditionally hunted pilot whales (n=14 males and n=13 females of different age groups) from the Faroe Islands. The main objectives of this study were to determine differences in the presence and concentrations of the compounds in the liver and plasma, how they depend on developmental stage (calves, sub adults, and adult females), and to assess maternal transfer of the compounds to suckling calves. Generally, the lipid weight (lw) concentrations of quantified POPs in the liver and plasma of pilot whales were positively correlated, and lw concentrations of most POPs did not differ between these matrices. However, concentrations of some individual POPs differed significantly (p0.05) between plasma and liver; CB-153 (p=0.044), CB-174 (p=0.027) and BDE-47 (p=0.017) were higher in plasma than in liver, whereas p,p'-DDE (p=0.004) and HCB (p0.001) were higher in liver than in plasma. POP concentrations differed between age/gender groups with lower levels in adult females than in juveniles. The relative distribution of compounds also differed between the age groups, due to the influence of the maternal transfer of the compounds. The results indicated that larger, more hydrophobic POPs were transferred to the offspring less efficiently than smaller or less lipid soluble compounds. Very low levels of both OH- and/or MeSO2-PCB and PBDE metabolites were found in all age groups, with no significant (p0.05) differences between the groups, strongly suggesting a very low metabolic capacity for their formation in pilot whales. The lack of difference in the metabolite concentrations between the age groups also indicates less maternal transfer of these contaminant groups compared to the precursor compounds.

Long-finned pilot whales (Globicephala melas) are rare visitors to the southern North Sea, but recently two individual strandings occurred on the Dutch coast. Both animals shared the same, unusual cause of death: asphyxiation from a common sole (Solea solea) stuck in their nasal cavity. This is a rare cause of death in cetaceans. Whilst asphyxiation has been reported in smaller odontocetes, there are no recent records of this occurring in Globicephala species. Here we report the stranding, necropsy and diet study results as well as discuss the unusual nature of this phenomenon. Flatfish are not a primary prey species for pilot whales and are rarely eaten by other cetaceans, such as harbour porpoises (Phocoena phocoena), in which there are several reports of asphyxiation due to airway obstruction by soles. This risk may be due to the fish’s flexible bodies which can enter small cavities either actively in an attempt to escape or passively due to the whale ‘coughing’ or ‘sneezing’ to rid itself of the blockage of the trachea. It is also possible that the fish enter the airways whilst the whale is re-articulating the larynx after trying to ingest large, oddly shaped prey. It is unlikely that the soles entered the airways after the death of the whales and we believe therefore that they are responsible for the death of these animals.

The primary bone pathology diagnoses recognized in cetacea are osteomyelitis and spondylosis deformans. In this study, we determined the prevalence, type, and severity of vertebral pathology in 52 pilot whales, a mass stranding species that stranded on Cape Cod, Massachusetts, between 1982 and 2000. Eleven whales (21%) had hyperostosis and ossification of tendon insertion points on and between vertebrae, chevron bones, and costovertebral joints, with multiple fused blocks of vertebrae. These lesions are typical of a group of interrelated diseases described in humans as spondyloarthropathies, specifically ankylosing spondylitis, which has not been fully described in cetacea. In severe cases, ankylosing spondylitis in humans can inhibit mobility. If the lesions described here negatively affect the overall health of the whale, these lesions may be a contributing factor in stranding of this highly sociable species.

The potential effects of exposing marine mammals to military sonar is a current concern. Dose-response relationships are useful for predicting potential environmental impacts of specific operations. To reveal behavioral response thresholds of exposure to sonar, we conducted 18 exposure/control approaches to 6 long-finned pilot whales. Source level and proximity of sonar transmitting one of two frequency bands (1-2kHz and 6-7kHz) were increased during exposure sessions. The 2-dimensional movement tracks were analyzed using a changepoint method to identify the avoidance response thresholds which were used to estimate dose-response relationships. No support for an effect of sonar frequency or previous exposures on the probability of response was found. Estimated response thresholds at which 50% of population show avoidance (SPLmax=170dB re 1μPa, SELcum=173dB re 1μPa2s) were higher than previously found for other cetaceans. The US Navy currently uses a generic dose-response relationship to predict the responses of cetaceans to naval active sonar, which has been found to underestimate behavioural impacts on killer whales and beaked whales. The navy curve appears to match more closely our results with long-finned pilot whales, though it might underestimate the probability of avoidance for pilot-whales at long distances from sonar sources. © 2014 Elsevier Ltd.

Numerous specimens of the 3 sibling species of the Anisakis simplex species complex (A. pegreffii, A. simplex (senso stricto)), and A. simplex sp. C) recovered from cetacean species stranded within the known geographical ranges of these nematodes were studied morphologically and genetically. The genetic characterization was performed on diagnostic allozymes and sequences analysis of nuclear (internal transcribed spacer [ITS] of ribosomal [r]DNA) and mitochondrial (mitochondrial [mt]DNA cox2 and rrnS) genes. These markers showed (1) the occurrence of sympatry of the 2 sibling species A. pegreffii and A. simplex sp. C in the same individual host, the pilot whale, Globicephala melas Traill, from New Zealand waters; (2) the identification of specimens of A. pegreffii in the striped dolphin, Stenella coeruleoalba (Meyen), from the Mediterranean Sea; and (3) the presence of A. simplex (s.s.) in the pilot whale and the minke whale, Balaenoptera acutorostrata Lacépède, from the northeastern Atlantic waters. No F1 hybrids were detected among the 3 species using the nuclear markers. The phylogenetic inference, obtained by maximum parsimony (MP) analysis of separate nuclear (ITS rDNA region), combined mitochondrial (mtDNA cox2 and rrnS) sequences datasets, and by concatenated analysis obtained at both MP and Bayesian inference (BI) of the sequences datasets at the 3 studied genes, resulted in a similar topology. They were congruent in depicting the existence of the 3 species as distinct phylogenetic lineages, and the tree topologies support the finding that A. simplex (s.s.), A. pegreffii, and A. berlandi n. sp. (= A. simplex sp. C) represent a monophyletic group. The morphological and morphometric analyses revealed the presence of morphological features that differed among the 3 biological species. Morphological analysis using principal component analysis, and Procrustes analysis, combining morphological and genetic datasets, showed the specimens clustering into 3 well-defined groups. Nomenclatural designation and formal description are given for A. simplex species C: the name Anisakis berlandi n. sp. is proposed. Key morphological diagnostic traits are as follows between A. berlandi n. sp. and A. simplex (s.s.): ventriculus length, tail shape, tail length/total body length ratio, and left spicule length/total body length ratio; between A. berlandi n. sp. and A. pegreffii: ventriculus length and plectane 1 width/plectane 3 width ratio; and between A. simplex (s.s.) and A. pegreffii: ventriculus length, left and right spicule length/total body length ratios, and tail length/total body length ratio. Ecological data pertaining to the geographical ranges and host distribution of the 3 species are updated.

Synchronous behavior, as a form of social interaction, has been widely reported for odontocete cetaceans observed at the sea surface. However, few studies have quantified synchronous behavior underwater. Using data from an animal-borne data recorder and camera, we described how a pair of deep-diving odontocetes, long-finned pilot whales, coordinated diving behavior. Diving data during overlapping periods of 3.7 h were obtained from two whales within a stable trio. The tagged whales made highly synchronous movements, and their dive durations differed only slightly (3 ± 3 s). The pair of whales maintained a constant and narrow vertical separation (ca. 3 m) throughout synchronous dives. The overall fluking rate for the same travel speed during synchronous dives was virtually the same as that during asynchronous dives, suggesting that synchronous behavior did not affect locomotion effort. In addition, a possible affiliative behavior was recorded by the animal-borne camera: another individual appeared in 8% of the frames, both with and without body contact to the tagged whale. The primary type of body contact was flipper-to-body. Our study, the first on underwater synchronous behavior and body contact of pilot whales, highlights the utility of using animal-borne devices for enabling new insights into social interactions.

Pollution is a threat to the health of marine mammals worldwide. Mass-strandings are poorly understood, but often involve pilot whales. However, there is limited information regarding pollution in long-finned pilot whales from Australia. Consequently, the profiles and levels of several pollutant classes were investigated in blubber of Tasmanian long-finned pilot whales. DDX levels were highest in all groups, followed by PCBs or MeO-PBDEs and lowest for PBDEs. The concentrations of all pollutants decreased with age in males. This is at least partly due to the growth dilution effect although it might also be caused by decreasing levels of PCBs, PBDEs, DDXs, HCB and CHLs in the environment. Fetus/mother ratios of higher chlorinated PCBs increased with the duration of pregnancy suggesting a preference for offloading via gestation rather than through lactation. Overall, the highest pollutant levels were found in the youngest animals.

This study was mainly funded by three naval organisations: the US Office of Naval Research, the Norwegian Ministry of Defense and the Netherlands Ministry of Defense. In addition, WWF-Norway, TOTAL Foundation and the Foundation Bleustein-Blanchet also contributed financially. Authors are employed by government (Norwegian Defense Research Establishment), independent no-profit (Netherlands Organization for Applied Scientific Research and Kelp Marine Research), or academic (University of St. Andrews) research organisations. No authors are employed by naval organisations. The funders had no role in study design, data analysis, or preparation of the manuscript. In cetaceans’ communities, interactions between individuals of different species are often observed in the wild. Yet, due to methodological and technical challenges very little is known about the mediation of these interactions and their effect on cetaceans’ behavior. Killer whales (Orcinus orca) are a highly vocal species and can be both food competitors and potential predators of many other cetaceans. Thus, the interception of their vocalizations by unintended cetacean receivers may be particularly important in mediating interspecific interactions. To address this hypothesis, we conducted playbacks of killer whale vocalizations recorded during herring-feeding activity to free-ranging long-finned pilot whales (Globicephala melas). Using a multi-sensor tag, we were able to track the whales and to monitor changes of their movements and social behavior in response to the playbacks. We demonstrated that the playback of killer whale sounds to pilot whales induced a clear increase in group size and a strong attraction of the animals towards the sound source. These findings provide the first experimental evidence that the interception of heterospecific vocalizations can mediate interactions between different cetacean species in previously unrecognized ways. Publisher PDF

CONSERVATION and management of natural systems often require biological research so that decisions can be based on scientific evidence ([Tracy and Brussard 1996][1]). Most biological research with application to conservation requires fieldwork, and fieldwork on animals involves a variety of

Background Lung cancer is the most commonly diagnosed neoplasm in humans, however this does not apply to other animal species. Living in an aquatic environment the respiratory system of cetaceans had to undergo unique adaptations in order to them to survive and cope with totally different respiratory pathogens and potentially carcinogens from those affecting humans. Case presentation This article discusses not only macroscopical, histopathological and immunohistochemical features of a pulmonary carcinoma with disseminated metastases in a long-finned pilot whale (Globicephala melas), as well as the immunohistochemical analysis performed on various tissues of cetaceans belonging to the genus Globicephala. On the necropsy examination of the carcass, multiple pulmonary nodules and generalised thoracic lymphadenomegaly were noted. Histologically, a malignant epithelial neoplasia was identified in the lung, thoracic lymph nodes, and adrenal gland. Immunohistochemical analysis revealed a pulmonary carcinoma. Vasculogenic mimicry and epithelial-to-mesenchymal transition phenotype, as suggested by cytomorphological and immunohistochemical characteristics, were observed. Conclusions A diagnosis of metastatic pulmonary carcinoma was determined, which to the author’s knowledge, appears to be not previously recorded in long-finned pilot whale species. This is also the first report of vasculogenic mimicry and epithelial-to-mesenchymal transition event in a spontaneous cancer from a cetacean species.

Visual species identification of cetacean strandings is difficult, especially when dead specimens are degraded and/or species are morphologically similar. The two recognised pilot whale species (Globicephala melas and Globicephala macrorhynchus) are sympatric in the North Atlantic Ocean. These species are very similar in external appearance and their morphometric characteristics partially overlap; thus visual identification is not always reliable. Genetic species identification ensures correct identification of specimens. Here we have employed one mitochondrial (D-Loop region) and eight nuclear loci (microsatellites) as genetic markers to identify six stranded pilot whales found in Galicia (Northwest Spain), one of them of ambiguous phenotype. DNA analyses yielded positive amplification of all loci and enabled species identification. Nuclear microsatellite DNA genotypes revealed mixed ancestry for one individual, identified as a post-F1 interspecific hybrid employing two different Bayesian methods. From the mitochondrial sequence the maternal species was Globicephala melas. This is the first hybrid documented between Globicephala melas and G. macrorhynchus, and the first post-F1 hybrid genetically identified between cetaceans, revealing interspecific genetic introgression in marine mammals. We propose to add nuclear loci to genetic databases for cetacean species identification in order to detect hybrid individuals., Sí