Liver and renal histopathology of North Atlantic long-finned pilot whales (Globicephala melas) contaminated with heavy metals and organochlorine compounds

Long-finned pilot whales (Globicephala melas) from the Faroe Islands are known to be heavily polluted with contaminants, such as mercury (Hg) and organochlorine compounds (OC). This is postulated to exert adverse health effects on whales as well as the human population who rely on its meat and blubber as food sources. It was therefore decided to conduct a screening pilot study to determine contaminant concentrations and histopathology of liver and renal tissues in a total of 14 specimens from this subpopulation. In blubber, the mean ∑OC concentration was 31,887 ng g^?1 lw (range: 18,170–47,425 ng g^?1 lw) of which ∑PCB concentration was 23,416 ng g^?1 lw (range: 13,947–34,543 ng g^?1 lw; n = 3). In liver, mean Hg concentration was 138 µg^?1 ww (range: 54–351 µg g^?1 ww; n = 7) and mean cadmium (Cd) concentration 15 µg g^?1 ww (range: 7–31 µg g^?1 ww; n = 7). In the kidney, the mean Cd concentration was 36 µg g^?1 ww (range: 23–47 µg g^?1 ww; n = 7). Of the liver Hg concentrations, two were at the suggested toxic threshold levels of 60 µg g^?1 ww and five were 2–6-fold above. Liver selenium (Se) : Hg was on average 1.11 (range: 0.97–1.41; n = 7) indicating that Se was in excess, which decreased the risk of acute Hg poisoning. Histopathological examinations showed high prevalence (>35%) of renal glomerular arteriosclerosis, glomerular capillary dilatation, dilatation and hyalinization of Bowman's space/capsule and tubular hyaline casts. In liver tissue, high prevalence was found for portal cell infiltrates, lipid granulomas, hepatocytic lipid accumulation, bile duct proliferation, lipid-filled Ito cells, and focal necrosis. In a single juvenile male, 4 of 8 renal and 3 of 7 liver lesions were present. Cadmium concentrations increased significantly in the presence of glomerular arteriosclerosis and a similar trend was found for tubular hyaline casts. Based on these findings, as well as the nature of the lesions, data indicate that the histopathological changes were a result of age and that contaminants are likely to be the co-factors in the development in at least three renal and four liver lesions.     

Average nanorough skin surface of the pilot whale (Globicephala melas, Delphinidae): considerations on the self-cleaning abilities based on nanoroughness

In aquatic environments, the biofouling process is assumed to initiate from the conditioning layer of absorbed organic carbon residues on wetted surfaces. Microfouling organisms attach to this conditioning layer, building up a biofilm on which further biofouling proceeds. In dolphins, biofouling reduces hydrodynamic efficiency and may negatively affect health if not managed. In the present study we examined the skin surface of the pilot whale (Globicephala melas). Employing cryo-scanning electron microscopic techniques combined with various sample preparations, the skin displayed an average nanorough surface characterized by a pattern of nanoridge-enclosed pores; the average pore size (approximately 0.20 µm²) was below the size of most marine biofouling organisms. Further, the implications of this type of surface to the self-cleaning abilities of the skin of pilot whales are discussed, based on reduced available space for biofouler attachment, the lack of any particular microniches as shelters for biofoulers, and the challenges of turbulent water flow and liquid-air interfaces during surfacing and jumping of the dolphin.     

A spatially explicit estimate of the prewhaling abundance of the endangered North Atlantic right whale

The North Atlantic right whale (NARW) (Eubalaena glacialis) is one of the world's most threatened whales. It came close to extinction after nearly a millennium of exploitation and currently persists as a population of only approximately 500 individuals. Setting appropriate conservation targets for this species requires an understanding of its historical population size, as a baseline for measuring levels of depletion and progress toward recovery. This is made difficult by the scarcity of records over this species’ long whaling history. We sought to estimate the preexploitation population size of the North Atlantic right whale and understand how this species was distributed across its range. We used a spatially explicit data set on historical catches of North Pacific right whales (NPRWs) (Eubalaena japonica) to model the relationship between right whale relative density and the environment during the summer feeding season. Assuming the 2 right whale species select similar environments, we projected this model to the North Atlantic to predict how the relative abundance of NARWs varied across their range. We calibrated these relative abundances with estimates of the NPRW total prewhaling population size to obtain high and low estimates for the overall NARW population size prior to exploitation. The model predicted 9,075–21,328 right whales in the North Atlantic. The current NARW population is thus <6% of the historical North Atlantic carrying capacity and has enormous potential for recovery. According to the model, in June–September NARWs concentrated in 2 main feeding areas: east of the Grand Banks of Newfoundland and in the Norwegian Sea. These 2 areas may become important in the future as feeding grounds and may already be used more regularly by this endangered species than is thought.     

Ecological tracers and at-sea observations document the foraging ecology of southern long-finned pilot whales (<Emphasis Type="Italic">Globicephala melas edwardii</Emphasis>) in Kerguelen waters

The food and feeding ecology of the poorly known southern long-finned pilot whale (Globicephala melas edwardii) was investigated using ecological tracers (muscle and skin δ¹³C and δ¹⁵N, and total mercury, Hg) on individuals from two mass strandings together with at-sea observations of live animals in Kerguelen waters, southern Indian Ocean. Sightings of cetaceans from longliners over 9 years (2003–2012) emphasized the regular occurrence of pilot whales in slope waters surrounding the archipelago. Tissue δ¹³C values (a proxy of consumer foraging habitat) suggest that pilot whales fed in slope waters and in oceanic subantarctic waters over the last months preceding stranding. Tissue δ¹⁵N values and Hg concentration (dietary proxies) indicate a high trophic position (~4.7) for the pilot whales, likely corresponding to a mixed diet of fish and squid (not crustaceans) of undetermined species. Both skin and muscle Hg concentrations were positively and linearly correlated to individual size with no concomitant δ¹⁵N changes, which can be interpreted as a progressive Hg accumulation in tissues of individuals throughout life with no parallel dietary shift. Skin and muscle Hg concentrations were linearly and positively related; hence, Hg skin could be used as a proxy of Hg concentration in muscle (a main Hg reservoir of the body). Kerguelen southern long-finned pilot whales were less Hg contaminated than most pilot whale populations studied so far, thus suggesting that they are not at a high risk to Hg-induced damages in the remote islands of the Southern Ocean.     

Tracking adult North Atlantic bluefin tuna (Thunnus thynnus) in the northwestern Atlantic using ultrasonic telemetry

Ultrasonic, depth-sensitive transmitters were used to track the horizontal and vertical movements, for up to 48 h, of 11 adult (136 to 340 kg estimated body mass) North Atlantic bluefin tuna (Thunnus thynnus Linnaeus). Fish were tracked in October 1995, September and October 1996, and August and September 1997 in the Gulf of Maine, northwestern Atlantic. The objective was to document the behavior of these fish and their schools in order to provide the spatial, temporal, and environmental information required for direct (i.e. fishery-independent) assessment of adult bluefin tuna abundance using aerial surveys. Transmitters were attached to free-swimming fish using a harpoon attachment technique, and all fish remained within the Gulf of Maine while being followed. Most of the bluefin tuna tagged on Stellwagen Bank or in Cape Cod Bay (and followed for at least 30 h) held a predominately easterly course with net horizontal displacements of up to 76 km d⁻¹. Mean (±SD) swimming depth for all fish was 14 ± 4.7 m and maximum depth for individuals ranged from 22 to 215 m. All but one fish made their deepest excursions, often single descents, at dawn and dusk. In general, adult bluefin tuna spent <8% of their time at the surface (0 to 1 m), <19% in the top 4 m, but >90% in the uppermost 30 m. Mean (±SD) speed over ground was 5.9 km h⁻¹, but for brief periods surpassed 20 to 31 km h⁻¹. Sea surface temperatures during tracking were 11.5 to 22.0 °C, and minimum temperatures encountered by the fish ranged from 6.0 to 9.0 °C. Tagged bluefin tuna and their schools frequented ocean fronts marked by mixed vertebrate feeding assemblages, which included sea birds, baleen whales, basking sharks, and other bluefin schools. Received: 19 July 1999 / Accepted: 25 March 2000     

Distribution of nutrients and AOU in the Mediterranean Water (North Atlantic Ocean)

Water samples were collected in the Northeast Atlantic Ocean (33-37^° N and 11-16^° W) in May 1983, and vertical profiles of chemical and physical parameters allowed us to identify the different water masses. A significant difference in salinity and its maximum can be found at different places and depths, as a consequence of the spreading of the Mediterranean Water (MW) from the Strait of Gibraltar. The apparent oxygen utilization shows a correlation with nutrients in North Atlantic Central Water. The relative percentages of water masses were assessed in order to examine the depth of maximum influence of MW as a function of latitude and to apply this knowledge to the black scabbard fish long-line strategy. The long-line fishing depth location was adapted to the specific MW distribution of the different banks with improvements in the mean catch efficiency. The thickness of the MW prevalence layer has a pronounced decrease to the south. MW influence clearly sinks and decrease to south-west. The highest percentage of the MW was found in the Gorringe Bank area and the lowest in the Madeira area.     

Population differentiation of the Atlantic spotted dolphin (Stenella frontalis) in the western North Atlantic, including the Gulf of Mexico

Information about the genetic population structure of the Atlantic spotted dolphin [Stenella frontalis (G. Cuvier 1829)] in the western North Atlantic would greatly improve conservation and management of this species in USA waters. To this end, mitochondrial control region sequences and five nuclear microsatellite loci were used to test for genetic differentiation of Atlantic spotted dolphins in the western North Atlantic, including the Gulf of Mexico (n=199). Skin tissue samples were collected from 1994–2000. Significant heterozygote deficiencies in three microsatellite loci within samples collected off the eastern USA coast prompted investigation of a possible Wahlund effect, resulting in evidence for previously unsuspected population subdivision in this region. In subsequent analyses including three putative populations, two in the western North Atlantic (n=38, n=85) and one in the Gulf of Mexico (n=76), significant genetic differentiation was detected for both nuclear DNA (R _ST=0.096, P≤0.0001) and mitochondrial DNA (Φ _ST=0.215, P≤0.0001), as well as for all pair-wise population comparisons for both markers. This genetic evidence for population differentiation coupled to known biogeographic transition zones at Cape Hatteras, North Carolina and Cape Canaveral, Florida, USA, evidence of female philopatry, and preliminary support for significant genetic differences between previously documented morphotypes of Atlantic spotted dolphins in coastal and offshore waters all indicate that the biology and life history of this species is more complex than previously assumed. Assumptions of large, panmictic populations might not be accurate in other areas where S. frontalis is continuously distributed (e.g., eastern Atlantic), and could have a detrimental effect on long-term viability and maintenance of genetic diversity in this species in regions where incidental human-induced mortality occurs.

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Diets of four deep-water scaphopod species (Mollusca) in the North Atlantic and the Nordic Seas

The buccal pouch contents of four deep-water scaphopod (Mollusca) species, Pulsellum affine, Pulsellum teres, Siphonodentalium lobatum and Polyschides olivi, collected in the North Atlantic and the Nordic Seas, were examined. The buccal pouches of the examined scaphopod species contained almost exclusively foraminifers, which agrees with previous studies on shallow and deep-water scaphopods. The scaphopods seemed to be engulfing at least some of the most common foraminifer species, which are distributed in the cold waters of the Nordic Seas or the temperate waters in the northernmost part of the North Atlantic. About half of the food species (30 foraminifer species) occurred in only one scaphopod species, nearly always in very low numbers, and seemed to be an unimportant food source. The remaining and more frequent food species occurred in two or more scaphopod species. A similarity tree was calculated with parsimony and presence–absence coding of the food species in the scaphopods. This resulted in the similarity tree: 100%{P. teres 79%[P. affine 70%(P. olivi and P. lobatum)]}, when branch support was measured with percentage jackknife stability. A general trend was found towards larger food items being consumed by larger scaphopod species, indicated by the positive correlation between the size of the scaphopods and the food species. It is suggested that microdistribution of the foraminifers and the large individual size of the foraminifers may be important factors in avoiding predation by scaphopods.     

A process-convolution approach to modelling temperatures in the North Atlantic Ocean

This paper develops a process-convolution approach for space-time modelling. With this approach, a dependent process is constructed by convolving a simple, perhaps independent, process. Since the convolution kernel may evolve over space and time, this approach lends itself to specifying models with non-stationary dependence structure. The model is motivated by an application from oceanography: estimation of the mean temperature field in the North Atlantic Ocean as a function of spatial location and time. The large amount of this data poses some difficulties; hence computational considerations weigh heavily in some modelling aspects. A Bayesian approach is taken here which relies on Markov chain Monte Carlo for exploring the posterior distribution.     

Continuous plankton records: The occurrence of apostome ciliates (Protozoa) on Euphausiacea in the North Atlantic Ocean and North Sea

The phoronts (resting cysts) of apostome ciliates have been recorded on Euphausia hemigibba, E. krohni, Meganyctiphanes norvegica, Nematoscelis megalops, Nyctiphanes couchi, Thysanoessa gregaria, T. inermis, T. longicaudata and T. raschi taken in the Continuous Plankton Recorder Survey of the North Atlantic Ocean and the North Sea. Variations in the percentage of host euphausiids infested by apostomes have been studied in relation to species, geography, body size and season. The percentage infestation was found to be related mainly to the body size of the hosts. The apostomes are believed to be species that develop either in the exuviae discarded exoskeletons or corpses of the euphausiids, or in the digestive systems of predators of the euphausiids.     

Molecular divergence between North Atlantic and Indo-West Pacific Cladophora albida (Cladophorales: Chlorophyta) isolates as indicated by DNA-DNA hybridization

Genomic relationships between North Atlantic, Australian and Japanese isolates of the benthic seaweed Cladophora albida (Huds.) Kütz. were examined in 1987 by means of DNA-DNA hybridization. The data indicate that C. albida can be divided into a North Atlantic and an Indo-West Pacific group with an intergroup hybridization response of 25 to 30% and 5.5° to 6.0°C for hybridization percentage and T _me the melting temperature reduction of hybridized sequences, respectively. This level of genome divergence is considerably higher than that observed in most other eukaryotes. The separation between the two C. albida groups presumably dates back to the closure of the Asian part of the Tethys Ocean, about 12 million years ago. The data also indicate that transatlantic C. albida populations have a greater genetic inter-relatedness than have Japanese and Australian populations. In C. albida there is no clear correlation between molecular evolution and the evolution of morphological traits. C. albida and C. rupestris (L.) Kütz have hardly any DNA sequences in common.     

Diet of the social groups of long-finned pilot whales (<Emphasis Type="Italic">Globicephala melas</Emphasis>) in the Strait of Gibraltar

The Strait of Gibraltar is inhabited throughout the year by a group of pilot whales (Globicephala melas), but their spatial distribution varies between Summer and Autumn. In this paper, we have used carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) stable isotope signatures to investigate the differences in diet amongst seasons, sex and stable social units. Skin samples were collected from 56 individually photo-identified pilot whales during Autumn 2005 and Summer 2006. These individuals were genetically sexed and their isotopic signature determined. The level of inter-individual association both within and between stable social units were compared to Euclidean distances between individual isotopes signatures. No differences in either δ¹⁵N or δ¹³C were found according to the sex of individuals, but significant seasonal differences were found in δ¹⁵N, although not in the δ¹³C values. This suggests that pilot whales are resident year round in the Strait, a finding supported by independent photo-identification. The variation in δ¹⁵N could reflect a shift in pilot whale diet through the year, with pilot whales feeding at a higher trophic level in Autumn compared to Summer. This could also represent a change in the diet of pilot whale prey species. The δ¹³C values were significantly different amongst the four stable social units sampled and individual δ¹³C values were significantly related to the level of inter-individual association, while no relationship was found for δ¹⁵N. These results suggest that within the same general area (i.e. the Strait of Gibraltar), there is some level of specialisation in habitat or prey choice between pilot whales social units.     

Phylogeography of the rock-pool copepod Tigriopus brevicornis (Harpacticoida) in the northern North Atlantic, and its relationship to other species of the genus

We investigated relationships among North Atlantic Tigriopus brevicornis populations and their relationships to Mediterranean T. fulvus and North American T. californicus, using crossing experiments and mitochondrial DNA sequencing. All T. brevicornis populations tested were interfertile, while interspecific crosses produced either no offspring or offspring that did not survive past the larval stage, with the exception of a few T. brevicornis × T. californicus crosses that produced mature adults. DNA sequencing of a fragment of mitochondrial cytochrome oxidase I (COI) showed that samples of T. brevicornis from Iceland, the Faroes, Ireland, Scotland and Nova Scotia formed a single shallow clade. In contrast, T. brevicornis from more southern populations in France and Portugal formed a clade with substantially greater branch lengths. Tigriopus brevicornis was monophyletic, and T. brevicornis plus Mediterranean T. fulvus were together also monophyletic. The phylogeography of T. brevicornis closely mirrored that found in T. californicus, with substantially reduced interpopulation divergence at northern latitudes. The known distribution of T. brevicornis in Iceland and the Faroes is shown and dispersal mechanisms and habitat selection briefly discussed.     

Bowhead whale (Balaena mysticetus) growth and feeding as estimated by δ13C techniques

Tissue ¹³C of bowhead whale, Balaena mysticetus, varies seasonally in response to geographic variations in the ¹³C of prey organisms consumed along the annual migratory route between the Bering, Chukchi and Beaufort Seas. Seasonal changes in body ¹³C of whales taken in 1986 from Alaskan waters provide a means of estimating energy intake from the different habitats in which the whales feed. Adult bowheads do not show significant seasonal shifts in the ¹³C of muscle and visceral fat. Their enrichment in ¹³C relative to subadult whales suggests that they acquire most of their food from fall and winter feeding or from unsampled parts of the summer range where zooplankton are enriched in ¹³C. Young individuals however, undergo marked seasonal shifts indicating that they feed heavily both in the summer and autumn or winter. Oscillations in ¹³C along the length of the baleen also provide a measure of age. Baleen growth rates decline as the whales age and provide a means of correcting for wear loss and allow aging of individuals less than 12 yr old. Although body length is a poor indicator of age in young bowhead whales, baleen length is closely correlated. Growth rates of bowheads after Year 1 are slow ( 0.4m yr^-1) and up to 20 yr are required to reach the assumed length of sexual maturity at 13 to 14m.     

Temporal separation of two fin whale call types across the eastern North Pacific

Fin whales (Balaenoptera physalus) produce a variety of low-frequency, short-duration, frequency-modulated calls. The differences in temporal patterns between two fin whale call types are described from long-term passive acoustic data collected intermittently between 2005 and 2011 at three locations across the eastern North Pacific: the Bering Sea, off Southern California, and in Canal de Ballenas in the northern Gulf of California. Fin whale calls were detected at all sites year-round, during all periods with recordings. At all three locations, 40-Hz calls peaked in June, preceding a peak in 20-Hz calls by 3–5 months. Monitoring both call types may provide a more accurate insight into the seasonal presence of fin whales across the eastern North Pacific than can be obtained from a single call type. The 40-Hz call may be associated with a foraging function, and temporal separation between 40- and 20-Hz calls may indicate the separation between predominately feeding behavior and other social interactions.     

Genetic differentiation in a pelagic crustacean (Meganyctiphanes norvegica: Euphausiacea) from the North East Atlantic and the Mediterranean Sea

Meganyctiphanes norvegica (M. Sars)is a pelagic crustacean that plays a key role in marine food webs of North Atlantic Ocean and marginal seas. We studied eight population samples collected in the European Atlantic and Mediterranean Sea. By means of single strand conformation polymorphism analysis (SSCP) and direct sequencing, we investigated a segment of 158 base pairs of the mitochondrial gene coding for the subunit 1 of NADH dehydrogenase. We found 12 sequence variants among the 385 individuals studied. Analysis of molecular variance (AMOVA) showed that 14.75% of the total genetic variability was explained by differences between populations, thus indicating absence of panmixia for these populations. Pairwise comparisons revealed three distinct genetic pools: the first one represented by Cadiz Bay, the second one by the Ligurian Sea, and the third one included all the NE Atlantic samples. We also investigated one population from the Alboran Sea (within the Mediterranean basin, east of the Strait of Gibraltar). This population was found to be genetically intermediate between the NE Atlantic samples and the Ligurian sample, suggesting that the restriction to the gene flow is not associated with the Strait of Gibraltar, but possibly with the Oran–Almeria oceanographic front. The present work indicates that M. norvegica, although endowed with a high dispersal capacity because of its pelagic habit, can develop separate breeding units inside the same oceanic basin (the Atlantic). Furthermore, the Ligurian sample should be considered as a distinct evolutionary entity, separated from the Atlantic population. Received: 2 May 1999 / Accepted: 26 November 1999     

Lunar rhythms of vertical migrations coded in otolith microstructure of North Atlantic lanternfishes,genus Hygophum (Myctophidae)

Otoliths of five Hygophum species were examined by means of light and scanning electron microscopy. In otoliths of four species (H. benoiti, H. macrochir, H. reinhardtii and H. taaningi) a strong cyclic pattern of the incremental structure was observed. In the fifth species (H. hygomii) such a pattern did not exist. An analysis of archival data on mesopelagic collections suggested three types of Hygophum spp. migratory behavior in relation to the lunar cycle which corresponded with the otolith microstructure. In H. hygomii only limited influence of moon phase on the uppermost range of night-vertical migration toward the surface was observed. The abundant nighttime occurrence of this species moved from the 0 to 50 m into the 50 to 100 m depth strata at full moon. In H. benoiti, a great part of the population, mainly juveniles, showed a tendency toward cessation of the vertical migrations during the first and fourth quarters of the lunar cycle. H. macrochir and H. taaningi had the strongest correlation of behavior with the lunar cycle. Both species showed arrested vertical migrations at the new moon phase, staying at day depths during the night, i. e., below 400 m. Thus, sequences of clear growth increments in otoliths represented a fast-growth period associated with the night migration to the warm surface layers, while bands without easily distinguishable incremental structure were interpreted as a period of slow growth in deep, cold waters due to limitation of the upward migration range occurring approximately at new moon.     

Flexible foraging movements of leatherback turtles across the North Atlantic Ocean

Some marine species have been shown to target foraging at particular hotspots of high prey abundance. However, we show here that in the year after a nesting season, female leatherback turtles (Dermochelys coriacea) in the Atlantic generally spend relatively little time in fixed hotspots, especially those with a surface signature revealed in satellite imagery, but rather tend to have a pattern of near continuous traveling. Associated with this traveling, distinct changes in dive behavior indicate that turtles constantly fine tune their foraging behavior and diel activity patterns in association with local conditions. Switches between nocturnal vs. diurnal activity are rare in the animal kingdom but may be essential for survival on a diet of gelatinous zooplankton where patches of high prey availability are rare. These results indicate that in their first year after nesting, leatherback turtles do not fit the general model of migration where responses to resources are suppressed during transit. However, their behavior may be different in their sabbatical years away from nesting beaches. Our results highlight the importance of whole-ocean fishing gear regulations to minimize turtle bycatch.     

Seasonal distribution of modern coccolithophores in the western North Atlantic Ocean

The seasonal successions of standing crop and floral assemblage were studied for oceanic coccolithophores at 5 weather stations in the western North Atlantic Ocean. The coccolithophores were less populous during winter at the northern stations, whereas they became scarce during summer at the southern stations. The average standing crop in the surface water was quite consistent throughout all climatic regions, being a few tens of thousand cells per liter. The average standing crop at the 100 m level showed a clear trend of increasing southwards. The population at this depth was only one-tenth as large as that of the surface level at the northernmost station, whereas it was almost two-thirds as large as the surface levels at the southernmost station. Many of the major species exhibited seasonally biased occurrences. The trend was mostly repeated in consecutive years, although the magnitude of their abundances fluctuated from year to year in some case. The average species composition was obtained for the 5 weather stations at the surfacewater (surface and 20 m levels combined) and 100 m levels; Emiliania huxleyi dominated the flora at all stations. The relationship between occurrences of major species and ambient water temperature was also studied. Although some species were stenothermal, the majority exhibited eurythermal characteristics; this accounts for the high specific diversity of the coccolithophore community throughout the wide range of waters studied.Contribution No. 2899 of the Lamont-Doherty Geological Observatory of Columbia University.