Holocene environmental changes and the seal (Phocidae) fauna of the Baltic Sea: coming,going and staying

• 1 The occurrence of different species of seal (Phocidae) during the Holocene in north‐eastern Europe was influenced by a changing geological situation over the last 12 000 years, characterized by alternating freshwater, brackish and marine stages and by spatially and temporally limited opportunities for migration of marine species into the Baltic basin. The patterns of immigration by ringed seal Phoca hispida, harp seal Phoca groenlandica, grey seal Halichoerus grypus and harbour seal Phoca vitulina are reviewed here in order to understand present distributions and population sizes. Furthermore, the future of their populations is analysed.
• 2 Phoca hispida immigrated into the Baltic basin during its brackish Yoldia Stage at the Late Glacial/Early Holocene boundary and has persisted in some disjunct populations until now. A second immigration followed at the beginning of the Littorina Transgression – which resulted in the present stage of the Baltic Sea, obviously caused by the strong cooling event that occurred 6200 calibrated ¹⁴C years BC. Its present status is not threatened, but global warming will restrict suitable habitat.
• 3 Phoca groenlandica had a mid‐Holocene intermezzo in the Baltic Sea. Its later disappearance was probably caused by hunting of subadult animals by humans.
• 4 Halichoerus grypus and Phoca vitulina also immigrated at the beginning of the present stage of the Baltic Sea. Whereas H. grypus has been common from this beginning point until modern times, the population of P. vitulina persisted at a low level for millennia. Only since the 18th century has its population increased and spread over the whole Baltic Sea.

     

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.     

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.     

A review of the world's Grey seal population

A review is made of current information on the numbers of Grey seal (Halichoerus grypus throughout its range. The world stock is estimated at 52,500 with 5000 in the north-west Atlantic, 42,500 in the north-east Atlantic (34,200 in Great Britain) and 5000 in the Baltic.     

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.     

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.     

The occurrence of Corynosoma strumosum in the grey seal,Halichoerus grypus,caught off the Atlantic coast of Ireland

The occurrence, location and sex ratio of Corynosoma strumosum (Acanthoceophala: Polymorphidae) from 26 juvenile grey seals, Halichoerus grypus, by-caught from around the Inishkea Islands, Co. Mayo, from March to June 1997 were recorded. The location of C. strumosum within the seal did not vary, with all worms being found in the small intestine. The prevalence of infection was 100%, with no other acanthocephalan species being recorded. The mean abundance was 416 (range 80-846) C. strumosum per seal. There was no significant change in abundance by month. The sex ratio of mature female to mature male parasites was 1.2:1. The lengths of female and male worms did not differ significantly.     

The occurrence of Mycoplasma phocicerebrale, Mycoplasma phocidae, and Mycoplasma phocirhinis in grey and common seals (Halichoerus grypus and Phoca vitulina) in the United Kingdom

Following the isolation of Mycoplasma phocicerebrale from the flipper wound of a grey seal (Halichoerus grypus) in Cornwall, UK, surveillance for Mycoplasma species was extended to include other seals rescued or found dead around the UK. Mycoplasma phocicerebrale was frequently detected from the teeth of seals and from infected wounds and respiratory tracts. Mycoplasma phocirhinis, Mycoplasma phocidae, and some unidentified Mycoplasma species were also detected. Mycoplasma phocicerebrale and M. phocidae were the only bacteria consistently identified from the wound infections, but their role in respiratory and other diseases remains unknown, as other bacteria were also isolated from respiratory sites.     

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.     

Patterns of parental relatedness and pup survival in the grey seal (Halichoerus grypus)

Levels of parental relatedness can affect offspring survival and susceptibility to disease. We investigated parental relatedness of live and dead Halichoerus grypus pups between and within island populations and between possible causes of mortality. Nine microsatellites were used to calculate internal relatedness (IR) and standardized mean d2. We find that pups with higher than average levels of IR have significantly lower survival and that this varied between island populations and that certain loci contributed to the effect more than others. Although, there were no significant differences between causes of mortality, peritonitis, infection and stillborn had the highest levels of IR. These results provide evidence that parental relatedness is an important determinant of pre-weaning pup survival in the grey seal and that this may vary with cause of mortality given a larger sample size.     

Exposing the grey seal as a major predator of harbour porpoises

Harbour porpoises (Phocoena phocoena) stranding in large numbers around the southern North Sea with fatal, sharp-edged mutilations have spurred controversy among scientists, the fishing industry and conservationists, whose views about the likely cause differ. The recent detection of grey seal (Halichoerus grypus) DNA in bite marks on three mutilated harbour porpoises, as well as direct observations of grey seal attacks on porpoises, have identified this seal species as a probable cause. Bite mark characteristics were assessed in a retrospective analysis of photographs of dead harbour porpoises that stranded between 2003 and 2013 (n = 1081) on the Dutch coastline. There were 271 animals that were sufficiently fresh to allow macroscopic assessment of grey seal-associated wounds with certainty. In 25% of these, bite and claw marks were identified that were consistent with the marks found on animals that had tested positive for grey seal DNA. Affected animals were mostly healthy juveniles that had a thick blubber layer and had recently fed. We conclude that the majority of the mutilated harbour porpoises were victims of grey seal attacks and that predation by this species is one of the main causes of death in harbour porpoises in The Netherlands. We provide a decision tree that will help in the identification of future cases of grey seal predation on porpoises.     

The respiratory system of Halarachne halichoeri (Halarachnidae: Gamasida: Anactinotrichida)

Halarachne halichoeri Allman (Halarachnidae: Anactinotrichida), a parasitic mite occuring in the nasal cavities of the grey seal, Halichoerus grypus , has an unusual apneustic respiratory system, which is described in detail. The main functions of this system are to facilitate gaseous (tracheal) respiration and prevent foreign material being forced into the tracheae under increased pressure experience when the host seal dives. Possible responses of the system to the rapidly changing pressure regimes are described.     

Archaeological evidence of former occurrence and changes in fishes, amphibians, birds, mammals and molluscs in the Wadden Sea area

Animal remains are well preserved in archaeological sites, especially the terp sites, of the Wadden Sea area of Denmark, Germany and The Netherlands. Here, we provide an overview on the wild mammals, birds, fishes, amphibians and molluscs found in coastal sites dating from 2700 to 2600 B.C. and 700 B.C. to A.D. 1600. Coastal people used a variety of animal species for food and other purposes. Hunting, fowling, fishing and agriculture did not have much influence on wild stocks in the period from the late Bronze Age/early Iron Age until the late Middle Ages. However, large changes to the landscape were made in the late Middle Ages by diking and damming. As a result, some species such as the northern vole (Microtus oeconomus) and the natterjack toad (Bufo calamita) disappeared from the area except for some dune districts on the islands, and others became rare, such as the grey seal (Halichoerus grypus) and the lagoon cockle (Cerastoderma lamarcki). New habitats arose for birds of dry meadows and fields, like lapwing (Vanellus vanellus) and black-tailed godwit (Limosa limosa). Sturgeon (Acipenser sturio) disappeared from the Wadden Sea within a few decades since A.D. 1890 due to the destruction of spawning grounds by damming and high exploitation pressure. Our findings are important for the ecological history of the region.     

Effects of environment on grey seal (Halichoerus grypus) pup mortality. Studies on the Isle of May

Breeding sites, including beach environment and topography, play a significant role in the causes of death of grey seal ( Halichoerus grypus ) pups, in particular the incidence of starvation, peritonitis/navel ill, other infectious diseases and injuries as a result of adult-pup interactions. The bacteria involved are opportunistic invaders. A wide and differing sex ratio in dead pups was found on different beaches and this has not been satisfactorily explained.     

Modelling regional differences in 'sealworm', Pseudoterranova decipiens (Nematoda, Ascaridoidea), infections in some North Atlantic cod, Gadus morhua, stocks

The ascaridoid nematode Pseudoterranova decipiens has a complex life-cycle. Adult worms reproduce in the stomach of grey seals, Halichoerus grypus , and other pinnipeds, and the larvae are found encysted in the musculature of Atlantic cod, a common host. An increase in infection levels in Canadian, Icelandic and Norwegian cod stocks over the last decade has been attributed to recent increases in seal population numbers. A simple model is proposed for the accumulation of larval parasites in the fish through its food. The model describes how regional differences in fish growth can also explain some differences in infection levels.     

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.     

Could Seals Prevent Cod Recovery in the Baltic Sea?

Fish populations are increasingly affected by multiple human and natural impacts including exploitation, eutrophication, habitat alteration and climate change. As a result many collapsed populations may have to recover in ecosystems whose structure and functioning differ from those in which they were formerly productive and supported sustainable fisheries. Here we investigate how a cod (Gadus morhua) population in the Baltic Sea whose biomass was reduced due to a combination of high exploitation and deteriorating environmental conditions might recover and develop in the 21st century in an ecosystem that likely will change due to both the already started recovery of a cod predator, the grey seal Halichoerus grypus, and projected climate impacts. Simulation modelling, assuming increased seal predation, fishing levels consistent with management plan targets and stable salinity, shows that the cod population could reach high levels well above the long-term average. Scenarios with similar seal and fishing levels but with 15% lower salinity suggest that the Baltic will still be able to support a cod population which can sustain a fishery, but biomass and yields will be lower. At present knowledge of cod and seal interactions, seal predation was found to have much lower impact on cod recovery, compared to the effects of exploitation and salinity. These results suggest that dual management objectives (recovery of both seal and cod populations) are realistic but success in achieving these goals will also depend on how climate change affects cod recruitment.     

Characterization of xenobiotic-metabolizing cytochrome P450 (CYP) forms in ringed and grey seals from the Baltic Sea and reference sites

Earlier studies have shown that members of the cytochrome P4501 (CYP1) enzyme family are constitutively expressed, and are elevated in the livers of ringed seals (Phoca hispida) and grey seals (Halichoerus grypus) living in the heavily polluted Baltic Sea. In this study, we compared the expression profiles of several additional CYP enzymes in the liver and extrahepatic tissues of Baltic ringed and grey seals with the corresponding CYP expression in seals from relatively unpolluted waters. We used marker enzyme activity levels, diagnostic inhibitors and immunoblot analysis to assess members of the CYP2A, CYP2B, CYP2C, CYP2D, CYP2E and CYP3A sub-families. Coumarin 7-hydroxylation (COH), a marker of CYP2A activity, was high in the liver and the lungs of all the studied seal populations. The presence of a putative CYP2A form in these seals was further supported by the strong inhibition of COH activity by a chemical inhibitor and by an anti-CYP2A5 antibody. However, antibodies to human and rodent CYP2B, CYP2C and CYP2E forms did not recognize any proteins in these seal species. Dextromethorphan O-demethylation (marker for CYP2D activity) and chlorzoxazone 6-hydroxylation (marker for CYP2E activity) were measurable in the livers of all the seals we studied. Both activities were elevated in the Baltic seal populations, showed a strong positive correlation with CYP1A activity and were at least partly inhibited by a typical CYP1A inhibitor, alpha-naphthoflavone. Further studies are needed to determine the presence and characteristics of CYP2D and CYP2E enzymes in ringed and grey seals. Testosterone 6beta-hydroxylation, a CYP3A marker, showed a relatively high level of activity in the livers of both seal species and was potently inhibited by ketoconazole, a CYP3A-selective inhibitor. The putative CYP3A activity showed an opposing geographical trend to that of CYP2D and CYP2E, since it was elevated in the control area. CYP3A protein levels, revealed by immunoblotting, showed a positive correlation with testosterone 6beta-hydroxylation. We conclude tentatively that CYP2A- and CYP3A-like enzymes are expressed in ringed and grey seals, but that CYP2B- and CYP2C-like ones are not. Further information on the individual contaminant profile is needed before any conclusions can be drawn on a possible connection between the varying CYP expressions and the contaminant load.