New polymorphic microsatellite markers for California sea lions (Zalophus californianus)

Nine microsatellite loci were isolated and characterized from California sea lions (Zalophus californianus). In addition, two of five loci tested from harbour seal (Phoca vitulina) produced a single, clear band in Z. californianus, as did one out of five loci from grey seal (Halichoerus grypus) and one out of two loci from elephant seal (Mirounga sp.). No locus tested from South American fur seal (Arctocephalus australis) amplified in Z. californianus. Locus variability was assessed in California sea lions from Los Islotes rookery, Baja California Sur, Mexico. All loci were variable, with allele numbers ranging from three to 12.     

Decreases in sealworm (Pseudoterranova sp. (p)) abundance in short-spined sea scorpion (Myoxocephalus scorpius scorpius) following declines in numbers of seals at Hvalseyjar, western Iceland

. Larval sealworm [Pseudoterranova sp. (p)] infections were surveyed in the short-spined sea scorpion (Myoxocephalus scorpius scorpius) in the shallow waters of the Hvalseyjar archipelago, western Iceland, in the summers of 1992, 1996 and 1999. Significant decreases in sealworm abundance in 1996 and 1999 were attributed to declines in local populations of grey (Halichoerus grypus) and common (Phoca vitulina) seals, final hosts for sealworm, between 1992 and 1998. The declines of the local seal populations were caused by over-exploitation, disturbances following hunting operation and accidental catch in fishing gear.     

Ten novel dinucleotide microsatellite loci cloned from the Galápagos sea lion (Zalophus californianus wollebaeki) are polymorphic in other pinniped species

We isolated and characterized 10 novel dinucleotide microsatellite loci from the Galápagos sea lion (Zalophus californianus wollebaeki) and tested their amplification utility in four further otariid species (Zalophus californianus californianus, Arctocephalus gazella, Arctocephalus australis and Eumetopias jubatus) and three phocid species (Hydrurga leptonyx, Halichoerus grypus and Phoca vitulina). All of the loci amplified polymorphic polymerase chain reaction (PCR) products in at least three species other than the Galápagos sea lion. These markers will be useful for studies of pinniped mating systems, genetic structure and genetic diversity.     

Kinematics of terrestrial locomotion in harbor seals and gray seals: Importance of spinal flexion by amphibious phocids

Pinnipeds are amphibious mammals with flippers, which function for both aquatic and terrestrial locomotion. Evolution of the flippers has placed constraints on the terrestrial locomotion of phocid seals. The detailed kinematics of terrestrial locomotion of gray (Halichoerus grypus) and harbor (Phoca vitulina) seals was studied in captivity and in the wild using video analysis. The seals exhibited dorsoventral undulations with the chest and pelvis serving as the main contact points. An anteriorly directed wave produced by spinal flexion aided in lifting the chest off the ground as the fore flippers were retracted to pull the body forward. The highest length‐specific speeds recorded were 1.02 BL/s for a gray seal in captivity and 1.38 BL/s for a harbor seal in the wild. The frequency and amplitude of spinal movement increased directly with speed, but the duty factor remained constant. Substrate did not influence the kinematics except for differences due to moving up or down slopes. The highly aquatic nature of phocids seals has restricted them to locomote on land primarily using spinal flexion, which can limit performance in speed and duration.     

The “Phoca standard”: An external molecular reference for calibrating recent evolutionary divergences

Comparison of the complete mitochondrial DNA (mtDNA) of the high-Arctic ringed seal (Phoca hispida) and the sub-Arctic harbour (P. vitulina) and grey (Halichoerus grypus) seals shows that they are genetically equidistant from one another. We relate the evolutionary divergence of the three species to expanding glaciation in the Arctic Basin and establish, in conjunction with mtDNA data, a standard reference for calibration of recent divergence events among mammalian taxa. In the present study, we apply the “Phoca standard” to the dating of divergences within the hominid phylogenetic tree. After determining the relative rates of substitution over all mitochondrial protein-coding genes in the different evolutionary lineages, we estimate that humans and chimpanzees diverged from each other 6.1 Mya (95% confidence limits: 5.2–6.9 Mya). The corresponding lower-limit divergence between common chimpanzee,Pan troglodytes, and pygmy chimpanzee,P. paniscus, occurred 3 (2.4–3.6) Mya, and the primary split within theP. troglodytes complex 1.6 (1.3–2.0) Mya. The analyses suggest that the split betweenGorilla andPan/Homo occurred 8.4 (7.3–9.4) Mya. They also suggest thatPongo (orangutan) and the lineage leading to gorillas, chimpanzees, and humans diverged 18.1 (16.5–19.6) Mya. The present analysis is independent of the hominid paleontological record and inferential morphological interpretations and thus is a novel approach to the lower-limit dating of recent divergences. Correspondence to: U. Arnason     

Global population structure and demographic history of the grey seal

Although the grey seal Halichoerus grypus is one of the most familiar and intensively studied of all pinniped species, its global population structure remains to be elucidated. Little is also known about how the species as a whole may have historically responded to climate‐driven changes in habitat availability and anthropogenic exploitation. We therefore analysed samples from over 1500 individuals collected from 22 colonies spanning the Western and Eastern Atlantic and the Baltic Sea regions, represented by 350 bp of the mitochondrial hypervariable region and up to nine microsatellites. Strong population structure was observed at both types of marker, and highly asymmetrical patterns of gene flow were also inferred, with the Orkney Islands being identified as a source of emigrants to other areas in the Eastern Atlantic. The Baltic and Eastern Atlantic regions were estimated to have diverged a little over 10 000 years ago, consistent with the last proposed isolation of the Baltic Sea. Approximate Bayesian computation also identified genetic signals consistent with postglacial population expansion across much of the species range, suggesting that grey seals are highly responsive to changes in habitat availability.     

Investigation of mercury concentrations in fur of phocid seals using stable isotopes as tracers of trophic levels and geographical regions

Recent studies have shown that the complementary analysis of mercury (Hg) concentrations and stable isotopic ratios of nitrogen (δ¹⁵N) and carbon (δ¹³C) can be useful for investigating the trophic influence on the Hg exposure and accumulation in marine top predators. In this study, we propose to evaluate the interspecies variability of Hg concentrations in phocids from polar areas and to compare Hg bioaccumulation between both hemispheres. Mercury concentrations, δ¹⁵N and δ¹³C were measured in fur from 85 individuals representing 7 phocidae species, a Ross seal (Ommatophoca rossii), Weddell seals (Leptonychotes weddellii), crabeater seals (Lobodon carcinophagus), harbour seals (Phoca vitulina), grey seals (Halichoerus grypus), ringed seals (Pusa hispida) and a bearded seal (Erignathus barbatus), from Greenland, Denmark and Antarctica. Our results showed a positive correlation between Hg concentrations and δ¹⁵N values among all individuals. Seals from the Northern ecosystems displayed greater Hg concentrations, δ¹⁵N and δ¹³C values than those from the Southern waters. Those geographical differences in Hg and stable isotopes values were likely due to higher environmental Hg concentrations and somewhat greater number of steps in Arctic food webs. Moreover, dissimilarities in feeding habits among species were shown through δ¹⁵N and δ¹³C analysis, resulting in an important interspecific variation in fur Hg concentrations. A trophic segregation was observed between crabeater seals and the other species, resulting from the very specific diet of krill of this species and leading to the lowest observed Hg concentrations.     

A Novel Tool to Mitigate By-Catch Mortality of Baltic Seals in Coastal Fyke Net Fishery

Developing methods to reduce the incidental catch of non-target species is important, as by-catch mortality poses threats especially to large aquatic predators. We examined the effectiveness of a novel device, a “seal sock”, in mitigating the by-catch mortality of seals in coastal fyke net fisheries in the Baltic Sea. The seal sock developed and tested in this study was a cylindrical net attached to the fyke net, allowing the seals access to the surface to breathe while trapped inside fishing gear. The number of dead and live seals caught in fyke nets without a seal sock (years 2008–2010) and with a sock (years 2011–2013) was recorded. The seals caught in fyke nets were mainly juveniles. Of ringed seals (Phoca hispida botnica) both sexes were equally represented, while of grey seals (Halichoerus grypus) the ratio was biased (71%) towards males. All the by-caught seals were dead in the fyke nets without a seal sock, whereas 70% of ringed seals and 11% of grey seals survived when the seal sock was used. The seal sock proved to be effective in reducing the by-catch mortality of ringed seals, but did not perform as well with grey seals.     

Improved estimates of digestion correction factors and passage rates for harbor seal (Phoca vitulina) prey in the northeast Atlantic

Diet composition in pinnipeds is widely estimated using hard prey remains recovered from feces. To estimate the size and number of prey represented in fecal samples accurately, digestion correction factors (DCFs) must be applied to measurements and counts of fish otoliths and cephalopod beaks. In this study, 101 whole prey feeding trials were conducted with six harbor seals (Phoca vitulina) and 18 prey species. We derived species‐ and grade‐specific estimates of digestion coefficients (DCs) and species‐specific recovery rates (RRs) to account for partial and complete digestion, respectively. Greater than 98% of otoliths were passed within three days of consumption. RRs were smallest for Atlantic salmon smolts (RR = 0.306, SE = 0.031) and increasingly larger for sandeels (RR = 0.494, SE = 0.017), flatfish (RR = 0.789, SE = 0.033), and large gadoids (RR = 0.944, SE = 0.034). Species‐specific otolith width DCs were smallest for Trisopterus species (DC = 1.14, SE = 0.015) and increasingly larger for flatfish (DC = 1.27, SE = 0.045), large gadoids (DC = 1.32, SE = 0.067) and sandeels (DC = 1.57, SE = 0.035). RRs were similar to those from gray seals (Halichoerus grypus), but harbor seal species‐ and grade‐specific DCs were generally smaller. Differences in partial and complete digestion rates among prey species and between seal species highlight the importance of applying DCFs when reconstructing diet.     

The identification of a new Giardia duodenalis assemblage in marine vertebrates and a preliminary analysis of G. duodenalis population biology in marine systems

Giardia duodenalis is an intestinal parasite of many vertebrates. The presence of G. duodenalis in the marine environment due to anthropogenic and wildlife activity is well documented, including the contributions from untreated sewage and storm water, agricultural run-off and droppings from terrestrial animals. Recently, studies have detected this protistan parasite in the faeces of marine vertebrates such as whales, dolphins, seals and shore birds. To explore the population biology of G. duodenalis in marine life, we determined the prevalence of G. duodenalis in two species of seal (Halichoerus grypus, Phoca vitulina vitulina and Phoca vitulina richardsi) from the east and west coasts of the USA, sequenced two loci from G. duodenalis-positive samples to assess molecular diversity and examined G. duodenalis distribution amongst these seals and other marine vertebrates along the east coast. We found a significant difference in the presence of G. duodenalis between east and west coast seal species. Only the zoonotic lineages of G. duodenalis, Assemblages A and B and a novel lineage, which we designated as Assemblage H, were identified in marine vertebrates. Assemblages A and B are broadly distributed geographically and show a lack of host specificity. Only grey seal (Halichoerus grypus) samples and one gull sample (Larus argentatus) from a northern location of Cape Cod, Massachusetts, USA, showed the presence of Assemblage H haplotypes; only one other study of harbour seals from the Puget Sound region of Washington, USA previously recorded the presence of an Assemblage H haplotype. Assemblage H sequences form a monophyletic clade that appears as divergent from the other seven Assemblages of G. duodenalis as these assemblages are from each other. The discovery of a previously uncharacterised lineage of G. duodenalis suggests that this parasite has more genetic diversity and perhaps a larger host range than previously believed.     

THE TROPHIC ROLE OF MARINE MAMMALS IN THE NORTHERN GULF OF ST. LAWRENCE

The trophic role of apex predators was evaluated in the northern Gulf of St. Lawrence ecosystem. An Ecopath model was developed for the period 1985–1987 prior to the collapse of commercially exploited demersal fish stocks in this area. Marine mammal trophic levels were estimated by the model at 4.1 for cetaceans, 4.4 for harp seals (Pagophilus groenlandicus), 4.7 for hooded seals (Cystophora cristata), 4.5 for gray seals (Halichoerus grypus), and 4.3 for harbor seals (Phoca vitulina). Harp seals were the third most important predator on vertebrate prey following large Atlantic cod (Gadus morhua) and redfish (Sebastes spp.). Different seal species preyed on different levels of the food chain. Harp seals preyed on most trophic groups, whereas larger seals, such as gray seals and hooded seals, mainly consumed higher trophic levels. The model suggested that apex predators had a negative effect on their dominant prey, the higher trophic level fish, but an indirect positive feedback on the prey of their preferred prey, mainly American plaice (Hippoglossoides platessoides), flounders, skates, and benthic invertebrates. Our results suggest that both marine mammals and fisheries had an impact on the trophic structure.     

Phocanema decipiens: Molting in seals

Infective larvae of the anisakine nematode Phocanema decipiens from the muscle of Atlantic cod (Gadus morhua) were fed to harbor seals (Phoca vitulina) and gray seals (Halichoerus grypus). During maturation in the stomach of seal hosts, P. decipiens molted twice; these molts are the third and fourth of its life cycle. The third molt occurred between the second and fifth days of infection. The third stage, i.e., infective larva entering the third molt, had a cuticular tooth ventral to the mouth; the fourth stage larva emerging from the third molt had three bilobed lips with dentigerous ridges. The fourth molt occurred between the 5th and 15th days in seals. A female nematode emerging from the fourth molt possesses a vulva and a vagina; a male possesses caudal alae, pre- and postanal papillae. Significant morphometric changes in nematodes were associated with both molts. Females and males of P. decipiens reached maturity after 15 to 25 days in seals. Ova were detected in the feces of the seal hosts as early as the 16th day.     

Seroprevalence of Antibodies against Seal Influenza A(H10N7) Virus in Harbor Seals and Gray Seals from the Netherlands

In the spring and summer 2014, an outbreak of seal influenza A(H10N7) virus infection occurred among harbor seals (Phoca vitulina) off the coasts of Sweden and Denmark. This virus subsequently spread to harbor seals off the coasts of Germany and the Netherlands. While thousands of seals were reported dead in Sweden, Denmark and Germany, only a limited number of seals were found dead in the Netherlands. To determine the extent of exposure of seals in the Netherlands to influenza A/H10N7 virus, we measured specific antibody titers in serum samples from live-captured seals and seals admitted for rehabilitation in the Netherlands by use of a hemagglutination inhibition assay and an ELISA. In harbor seals in 2015, antibodies against seal influenza A(H10N7) virus were detected in 41% (32 out of 78) pups, 10% (5 out of 52) weaners, and 58% (7 out of 12) subadults or adults. In gray seals (Halichoerus grypus) in 2015, specific antibodies were not found in the pups (n = 26), but in 26% (5 out of 19) of the older animals. These findings indicate that, despite apparent low mortality, infection with seal influenza A(H10N7) virus was geographically widespread and also occurred in grey seals.     

Examination of relationships between stable isotopes and cortisol concentrations along the length of phocid whiskers

Alaskan seals are found in remote and sometimes inaccessible locations, making it difficult to collect time‐series information. This study explores a novel method to examine temporal changes in diet and physiological status of ringed (Pusa hispida), spotted (Phoca largha), and harbor (Phoca vitulina) seals using cortisol concentrations and δ¹⁵N and δ¹³C stable isotopes (SIs) measured in serial sections of whiskers. As whiskers grow, whisker tissue is deposited sequentially making these measurements temporally aligned. Whisker cortisol presented in a distinct pattern with elevated concentrations at the root section followed by a curvilinear decline moving toward the tip of most whiskers. Comparing SIs at the root to the rest of the whiskers, δ¹³C values were slightly lower in ringed and harbor seal whiskers and δ¹⁵N values were slightly higher in harbor seal whiskers. The data were modeled controlling for the observed trends in cortisol concentrations and further associations between cortisol concentrations and SIs were detected in spotted and harbor seal whiskers. Additional research examining the source and stability of whisker cortisol is warranted. However, the methods presented here demonstrate that whiskers could prove valuable to gather long‐term and naturally aligned dietary and physiological information.     

Distribution of the Harp Seal Whelping Grounds in the White Sea 1960s–1990s

Analysis of air survey data has shown that from the 1960s to the 1990s the whelping grounds of the harp seal Phoca groenlandica formed at the end of February through the first days of March in two regions of the White Sea: in the eastern part of Bassein and in Gorlo. The areas of the whelping grounds varied depending on the current ice conditions. It was recorded that, despite ice conditions unfavorable for the pups' survival that developed in some years, in general the ice regime of the White Sea was favorable for reproduction of the White Sea population of the harp seal.     

Increasing numbers of harbour seals and grey seals in the Solent

Harbour seals (Phoca vitulina) and grey seals (Halichoerus grypus) both occur within the UK, but display regional contrasting population trends. While grey seals are typically increasing in number, harbour seals have shown varying trends in recent decades following repeated pandemics. There is a need for monitoring of regional and local populations to understand overall trends. This study utilized a 20‐year dataset of seal counts from two neighboring harbours in the Solent region of south England. Generalized additive models showed a significant increase in the numbers of harbour (mean 5.3–30.5) and grey (mean 0–12.0) seals utilizing Chichester Harbour. Conversely, in Langstone Harbour there has been a slight decrease in the number of harbour seals (mean 5.3–4.0). Accompanying photographic data from 2016 to 18 supports the increase in seal numbers within Chichester Harbour, with a total of 68 harbour and 8 grey seals identified. These data also show evidence of site fidelity of harbour seals in this area, with almost a quarter of animals resighted within the past three years. Overall, this long‐term study indicates an increasing number of both harbour and grey seals within the Solent. However, more research is required to identify the drivers of this trend.     

Meals on Wheels? A Decade of Megafaunal Visual and Acoustic Observations from Offshore Oil & Gas Rigs and Platforms in the North and Irish Seas

A decade of visual and acoustic detections of marine megafauna around offshore Oil & Gas (O&G) installations in the North and Irish Seas are presented. Marine megafauna activity was monitored visually and acoustically by Joint Nature Conservation Committee (JNCC) qualified and experienced Marine Mammal Observers (MMO) and Passive Acoustic Monitoring (PAM) Operators respectively, with real-time towed PAM in combination with industry standard software, PAMGuard. Monitoring was performed during routine O&G industrial operations for underwater noise mitigation purposes, and to ensure adherence to regulatory guidelines. Incidental sightings by off-effort MMOs and installation crew were also reported. Visual and acoustic monitoring spanned 55 non-consecutive days between 2004 and 2014. A total of 47 marine mammal sightings were recorded by MMOs on dedicated watch, and 10 incidental sightings of marine megafauna were reported over 10 years. Species included: harbour porpoise (Phocoena phocoena), Atlantic white-sided dolphin (Lagenorhynchus acutus), white beaked dolphin (Lagenorhynchus albirostris), common dolphin (Delphinus delphis), minke whale (Balaenoptera acutorostrata), common seal (Phoca vitulina), grey seal (Halichoerus grypus) and, basking shark (Cetorhinus maximus). Passive Acoustic Monitoring was conducted on two occasions in 2014; 160 PAM hours over 12 days recorded a total of 308 individual clicks identified as harbour porpoises. These appear to be the first such acoustic detections obtained from a North Sea drilling rig whilst using a typically configured hydrophone array designed for towing in combination with real-time PAMGuard software. This study provides evidence that marine megafauna are present around mobile and stationary offshore O&G installations during routine operational activities. On this basis, Environmental Impact Assessments (EIAs) for decommissioning O&G platforms should be carried-out on a case-by-case basis, and must include provisions for hitherto overlooked marine megafauna.     

Trichinella nativa in grey seal Halichoerus grypus: spill-over from a highly endemic terrestrial ecosystem

Trichinella spp. infections of marine mammals pose a human health risk in Arctic regions where game meat is an important part of the diet. In the last decade, the grey seal population of the sub-arctic Baltic Sea has reached a level at which hunting is possible and seal meat is now available for food. We conducted a survey on the occurrence of Trichinella spp. in grey seals Halichoerus grypus (n = 171) and ringed seals Phoca hispida botnica (n = 56) of the Baltic Sea in the coastal waters of Finland, a highly Trichinella sp. endemic area. Muscle samples were examined by a mechanically assisted digestion method during 2006-2010. One grey seal was positive for Trichinella nativa , while all samples from ringed seals were negative. Even though just 1 grey seal was infected, the finding here emphasizes the importance of proper meat inspection of seals intended for human consumption, especially in areas with high infection pressure.     

Rapid increase of the grey seal (<Emphasis Type="Italic">Halichoerus grypus</Emphasis>) breeding stock at Helgoland

Over the past 25 years, the grey seal (Halichoerus grypus) population in the North Sea increased on average at 5% per year, while founding a number of new colonies. Based on published data, this note compares recent trends in pup production at nine breeding sites, with focus on the German island of Helgoland. Since 1997, numbers born at Helgoland increased at 35% per year (95% CI: 29–42), reaching an all-time high of 53 in 2007. Yet, this was less than 1% of all grey seal offspring in the North Sea area. Some other small stocks had similarly high growth rates of 20–50%, which must be explained by immigration from the larger colonies, as sustained autochthonous increase cannot exceed 13% per year in the species. Further, the Helgoland breeding stock increased significantly faster than the neighbouring colonies at Amrum (13%) and in the Dutch Wadden Sea (20%), indicating higher offspring survival and/or higher relative influx of migrants. This is attributed to the fact that at Helgoland newborn seals are safe from floodwater. In the other two sites, in contrast, pups are born on sandbanks and are thus often hit by winterly storm tides, which poses a risk to their survival. Because of the favourable breeding conditions at Helgoland, this new colony is expected to soon become an important stronghold of the grey seal in the Dutch-German North Sea area.