Environmental drivers of population-level variation in the migratory and diving ontogeny of an Arctic top predator.

The development of migratory strategies that enable juveniles to survive to sexual maturity is critical for species that exploit seasonal niches. For animals that forage via breath-hold diving, this requires a combination of both physiological and foraging skill development. Here, we assess how migratory and dive behaviour develop over the first year of life for a migratory Arctic top predator, the harp seal Pagophilus groenlandicus, tracked using animal-borne satellite relay data loggers. We reveal similarities in migratory movements and differences in diving behaviour between 38 juveniles tracked from the Greenland Sea and Northwest Atlantic breeding populations. In both regions, periods of resident and transitory behaviour during migration were associated with proxies for food availability: sea ice concentration and bathymetric depth. However, while ontogenetic development of dive behaviour was similar for both populations of juveniles over the first 25 days, after this time Greenland Sea animals performed shorter and shallower dives and were more closely associated with sea ice than Northwest Atlantic animals. Together, these results highlight the role of both intrinsic and extrinsic factors in shaping early life behaviour. Variation in the environmental conditions experienced during early life may shape how different populations respond to the rapid changes occurring in the Arctic ocean ecosystem.

First record of plastic debris in the stomach of a hooded seal pup from the Greenland Sea.

Plastic debris is globally found around the world and the remote Arctic is no exception. Arctic true seals are sentinel species of marine pollution and represent the link between marine food webs and Arctic apex predators like polar bears and humans. With regard to true seals, ingested macroplastics have never been reported in an Arctic species. We harvested 10 harp seals Pagophilus groenlandicus and 8 hooded seals Cystophora cristata from the breeding grounds in the pack ice of the Greenland Sea. The digestive tract was inspected exclusively for the presence of macroplastics (>5 mm). Two pieces of single-use plastic were found in the stomach of a weaned hooded seal pup. This study indicates that young Arctic marine predators may ingest macroplastics, and therefore may be at risk during their early stages of life due to human caused plastic pollution even in the remote Arctic pack ice.

Pelagic vs Coastal-Key Drivers of Pollutant Levels in Barents Sea Polar Bears with Contrasted Space-Use Strategies.

In the Barents Sea, pelagic and coastal polar bears are facing various ecological challenges that may explain the difference in their pollutant levels. We measured polychlorinated biphenyls, organochlorine pesticides, polybrominated diphenyl ethers in fat, and perfluoroalkyl substances in plasma in pelagic and coastal adult female polar bears with similar body condition. We studied polar bear feeding habits with bulk stable isotope ratios of carbon and nitrogen. Nitrogen isotopes of amino acids were used to investigate their trophic position. We studied energy expenditure by estimating field metabolic rate using telemetry data. Annual home range size was determined, and spatial gradients in pollutants were explored using latitude and longitude centroid positions of polar bears. Pollutant levels were measured in harp seals from the Greenland Sea and White Sea-Barents Sea as a proxy for a West-East gradient of pollutants in polar bear prey. We showed that pelagic bears had higher pollutant loads than coastal bears because (1) they feed on a higher proportion of marine and higher trophic level prey, (2) they have higher energy requirements and higher prey consumption, (3) they forage in the marginal ice zones, and (4) they feed on prey located closer to pollutant emission sources/transport pathways.

Seroprevalence for Brucella spp. in Baltic ringed seals (Phoca hispida) and East Greenland harp (Pagophilus groenlandicus) and hooded (Cystophora cristata) seals.

Zoonotic infections transmitted from marine mammals to humans in the Baltic and European Arctic are of unknown significance, despite given considerable potential for transmission due to local hunt. Here we present results of an initial screening for Brucella spp. in Arctic and Baltic seal species. Baltic ringed seals (Pusa hispida, n = 12) sampled in October 2015 and Greenland Sea harp seals (Pagophilus groenlandicus, n = 6) and hooded seals (Cystophora cristata, n = 3) sampled in March 2015 were serologically analysed for antibodies against Brucella spp. The serological analyses were performed using the Rose Bengal Test (RBT) followed by a confirmatory testing of RBT-positive samples by a competitive-enzyme linked immunosorbent assay (C-ELISA). Two of the Baltic ringed seals (a juvenile male and a juvenile female) were seropositive thus indicating previous exposure to a Brucella spp. The findings indicate that ringed seals in the Baltic ecosystem may be exposed to and possibly infected by Brucella spp. No seropositive individuals were detected among the Greenland harp and hooded seals. Although our initial screening shows a zoonotic hazard to Baltic locals, a more in-depth epidemiological investigation is needed in order to determine the human risk associated with this.

Seasonal changes in background levels of deuterium and oxygen-18 prove water drinking by harp seals, which affects the use of the doubly labelled water method.

The aim of this study was to monitor seasonal changes in stable isotopes of pool freshwater and harp seal (Phoca groenlandica) body water, and to study whether these potential seasonal changes might bias results obtained using the doubly labelled water (DLW) method when measuring energy expenditure in animals with access to freshwater. Seasonal changes in the background levels of deuterium and oxygen-18 in the body water of four captive harp seals and in the freshwater pool in which they were kept were measured over a time period of 1 year. The seals were offered daily amounts of capelin and kept under a seasonal photoperiod of 69°N. Large seasonal variations of deuterium and oxygen-18 in the pool water were measured, and the isotope abundance in the body water showed similar seasonal changes to the pool water. This shows that the seals were continuously equilibrating with the surrounding water as a result of significant daily water drinking. Variations in background levels of deuterium and oxygen-18 in freshwater sources may be due to seasonal changes in physical processes such as precipitation and evaporation that cause fractionation of isotopes. Rapid and abrupt changes in the background levels of deuterium and oxygen-18 may complicate calculation of energy expenditure by use of the DLW method. It is therefore strongly recommended that analysis of seasonal changes in background levels of isotopes is performed before the DLW method is applied on (free-ranging) animals, and to use a control group in order to correct for changes in background levels.

Effects of Polar Bear and Killer Whale Derived Contaminant Cocktails on Marine Mammal Immunity.

Most controlled toxicity studies use single chemical exposures that do not represent the real world situation of complex mixtures of known and unknown natural and anthropogenic substances. In the present study, complex contaminant cocktails derived from the blubber of polar bears (PB; Ursus maritimus) and killer whales (KW; Orcinus orca) were used for in vitro concentration-response experiments with PB, cetacean and seal spp. immune cells to evaluate the effect of realistic contaminant mixtures on various immune functions. Cytotoxic effects of the PB cocktail occurred at lower concentrations than the KW cocktail (1 vs 16 µg/mL), likely due to differences in contaminant profiles in the mixtures derived from the adipose of each species. Similarly, significant reduction of lymphocyte proliferation occurred at much lower exposures in the PB cocktail (EC50: 0.94 vs 6.06 µg/mL; P < 0.01), whereas the KW cocktail caused a much faster decline in proliferation (slope: 2.9 vs 1.7; P = 0.04). Only the KW cocktail modulated natural killer (NK) cell activity and neutrophil and monocyte phagocytosis in a concentration- and species-dependent manner. No clear sensitivity differences emerged when comparing cetaceans, seals and PB. Our results showing lower effect levels for complex mixtures relative to single compounds suggest that previous risk assessments underestimate the effects of real world contaminant exposure on immunity. Our results using blubber-derived contaminant cocktails add realism to in vitro exposure experiments and confirm the immunotoxic risk marine mammals face from exposure to complex mixtures of environmental contaminants.

Hooded seal (Cystophora cristata) pups ingest snow and seawater during their post-weaning fast.

The purpose of this study was to evaluate the importance of exogenous water intake (snow/seawater) in hooded seal (Cystophora cristata) pups during their post-weaning fast. In this study, five hooded seal pups had ad lib access to snow and seawater for the first 12 and last 21 days of their post-weaning fast, respectively. Total body water and water flux were determined during both exposure periods by use of the tritiated water method. Blood samples were collected to monitor changes in hematocrit, plasma urea and plasma osmolality. Body mass loss was on average 0.36 kg day-1. Average total body water changed from 15.7 to 11.4 L, while total water influx changed from 15 to 18 mL day-1 kg-1 during snow and seawater exposure, respectively. Of this influx an average of 35% can be attributed to metabolic water, while approximately 8% was due to respiratory water influx. Interestingly, 56 and 58% of the total water influx was due to snow and seawater ingestion, respectively, amounting to 8 mL day-1 kg-1 snow (counted as liquid water) and 10 mL day-1 kg-1 seawater. Based on the results of the plasma parameters it is concluded that fasting hooded seal pups maintain water balance and homeostasis when access to snow or seawater is permitted. It is further concluded that snow and seawater intake, in addition to metabolic and respiratory water, is important for maintenance of water balance and excretion of urea during the post-weaning fast of hooded seal pups.

Serum chemistry and antibodies against pathogens in antarctic fur seals, Weddell seals, crabeater seals, and Ross seals.

Information on health parameters, such as antibody prevalences and serum chemistry that can reveal exposure to pathogens, disease, and abnormal physiologic conditions, is scarce for Antarctic seal species. Serum samples from Antarctic fur seals (Arctocephalus gazella, n=88) from Bouvetøya (2000-2001 and 2001-2002), and from Weddell seals (Leptonychotes weddellii, n=20), Ross seals (Ommatophoca rossii, n=20), and crabeater seals (Lobodon carcinophagus, n=9) from the pack-ice off Queen Maud Land, Antarctica (2001) were analyzed for enzyme activity, and concentrations of protein, metabolites, minerals, and cortisol. Adult Antarctic fur seal males had elevated levels of total protein (range 64-99 g/l) compared to adult females and pups (range 52-79 g/l). Antarctic fur seals had higher enzyme activities of creatine kinase, lactate dehydrogenase, and amylase, compared to Weddell, Ross, and crabeater seals. Antibodies against Brucella spp. were detected in Weddell seals (37%), Ross seals (5%), and crabeater seals (11%), but not in Antarctic fur seals. Antibodies against phocine herpesvirus 1 were detected in all species examined (Antarctic fur seals, 58%; Weddell seals, 100%; Ross seals, 15%; and crabeater seals, 44%). No antibodies against Trichinella spp., Toxoplasma, or phocine distemper virus (PDV) were detected (Antarctic fur seals were not tested for PDV antibodies). Antarctic seals are challenged by reduced sea ice and increasing temperatures due to climate change, and increased anthropogenic activity can introduce new pathogens to these vulnerable ecosystems and represent a threat for these animals. Our data provide a baseline for future monitoring of health parameters of these Antarctic seal species, for tracking the impact of environmental, climatic, and anthropogenic changes in Antarctica over time.

Application of the two-sample doubly labelled water method alters behaviour and affects estimates of energy expenditure in black-legged kittiwakes.

Despite the widespread use of the doubly labelled water (DLW) method in energetic studies of free-ranging animals, effects of the method on study animals are rarely assessed. We studied behavioural effects of two alternative DLW protocols. During two consecutive breeding seasons, 42 parent black-legged kittiwakes received either the commonly used two-sample (TS) or the less invasive single-sample (SS) DLW treatment. A third group served as a non-treated control. We evaluated the effect of treatment with respect to the time birds took to return to their nest after treatment and recaptures, and the nest attendance during DLW measurement periods. We found that TS kittiwakes took on average 20 times longer to return to their nest than SS kittiwakes after initial treatment, and nest attendance was reduced by about 40% relative to control birds. In contrast, nest attendance did not differ between control and SS kittiwakes. Estimates of energy expenditure of SS kittiwakes exceeded those of TS kittiwakes by 15%. This difference was probably caused by TS birds remaining inactive for extended time periods while at sea. Our results demonstrate that the common assumption that the TS DLW method has little impact on the behaviour of study subjects is in some circumstances fallacious. Estimates of energy expenditure derived by the SS approach may thus more accurately reflect unbiased rates of energy expenditure. However, the choice of protocol may be a trade-off between their impact on behaviour, and hence accuracy, and their differences in precision. Adopting procedures that minimize the impact of TS protocols may be useful.

Seawater drinking restores water balance in dehydrated harp seals.

The purpose of this study was to answer the question of whether dehydrated harp seals (Phoca groenlandica) are able to obtain a net gain of water from the intake of seawater. Following 24 h of fasting, three subadult female harp seals were dehydrated by intravenous administration of the osmotic diuretic, mannitol. After another 24 h of fasting, the seals were given 1,000 ml seawater via a stomach tube. Urine and blood were collected for measurement of osmolality and osmolytes, while total body water (TBW) was determined by injections of tritiated water. In all seals, the maximum urinary concentrations of Na(+) and Cl(-) were higher than in seawater, reaching 540 and 620 mM, respectively, compared to 444 and 535 mM in seawater. In another experiment, the seals were given ad lib access to seawater for 48 h after mannitol-induced hyper-osmotic dehydration. In animals without access to seawater, the mean blood osmolality increased from 331 to 363 mOsm kg(-1) during dehydration. In contrast, the blood osmolality, hematocrit and TBW returned to normal when the seals were permitted ad lib access to seawater after dehydration. In conclusion, this study shows that harp seals have the capacity to gain net water from mariposa (voluntarily drinking seawater) and are able to restore water balance after profound dehydration by drinking seawater.

Blood is a poor substrate for monitoring pollution burdens in phocid seals.

The present study investigated how concentrations of different organochlorines (OCs) vary with nutritional condition in adult harp seal (Phoca groenlandica) females in a captive fasting experiment and in the wild. During the first part of this study seals in good condition (N= 5) were fasted for a period of 28 days and blood and blubber samples were collected at days 1, 14 and 28. Blubber OC concentrations remained unchanged throughout the experimental period, and were consistently significantly higher than concentrations in the blood. In contrast to blubber OC levels, blood OC levels showed a significant time-dependant increase. During the fasting experiment the seals lost an average of 24 kg of body mass, which is less than half the natural annual variation in this species in the wild. The second part of this study compared OC concentrations in blood and blubber from seals collected at prime condition before the breeding season (N = 10) with animals collected during molt when condition is poor (N = 7). The average mass difference between the two groups was more than 40 kg. Blood levels of most OCs were significantly higher in the thin seals compared with the levels found in the fat seals. These differences in blood OC concentrations were much greater than what was found during the fasting experiment. For example sigmaPCB levels in the blood during the fasting experiment increased by approximately 83%, while the corresponding increase in blood levels of sigmaPCB between the two other samples was 720% (from 201 to 1,447 ng/g lipid). Blubber levels of OCs from the animals in the wild were significantly higher in the thin seals compared with the fat animals. In addition, the blubber levels of OCs were significantly higher than blood levels of OCs only for the fat seals. This study demonstrates the extreme variability present in the concentrations of OCs in blood of seals in response to change in condition. Since the natural variation in condition is extreme during phocid seals' annual cycles, we recommend that blood should not be used in studies of OCs where the aim of the study is to monitor OC levels for comparative purposes or time-trend analysis.