Since 2000, there has been a marked decline in the number of harbour seals in some regions around Britain; one possible contributing cause is competition for prey with sympatric grey seals. To explore one important aspect of this interaction, in this thesis the diet of harbour seals is estimated using analysis of hard prey remains recovered from faeces and compared with equivalent results for grey seals. To estimate coefficients to account for partial and complete digestion of hard prey remains, 100 whole prey feeding trials were conducted with six harbour seals and 18 prey species. Differences were found among prey species and between harbour and grey seals highlighting the importance of applying predator- and prey-specific digestion correction factors when reconstructing diet. In a comprehensive exploration of the diet of harbour seals around Britain, sandeel and flatfish dominated in the North Sea and large gadoids dominated on the Scottish west coast with seasonal pulses of pelagic prey. Variation in diet was linked to regional and seasonal differences in prey distribution and abundance. Sex-specific variation in harbour seal diet was examined in four regions. The main difference detected was in The Wash, where female diet quality was significantly higher than males in winter, which appeared to be driven by greater consumption of pelagic prey by female seals associated with seasonal energetic requirements of their annual life cycle. Comparison of the diet of harbour and grey seals revealed regional differences in diet composition, diversity and quality between the two species. However, there was no consistent pattern in this variation in relation to regional variation in harbour and grey seals population trajectories and no clear evidence for interspecific competition for prey. Future work should focus on an integrated investigation of prey abundance and distribution, and seal diet and foraging behaviour/distribution.

Animal communication systems are complex, but in many species relatively little is known about how they develop. Insight into the development of adult repertoires, considering how factors such as genetics, environment and learning contribute to signal changes, provides a more comprehensive understanding of communication. This thesis documented vocal learning and development in the grey seal. In chapter 3, vocal repertoires of grey seal pups were recorded from birth through their first year, and were compared to calls of other seals across life stages, from pups to adults. By examining call parameters that were similar in animals of the same age, size, and sex, changes that may be attributed to physical development and morphology were identified. Past studies have indicated that seals may also have advanced vocal learning abilities, however the extent of these capabilities was unknown. In this thesis, vocal learning in juvenile grey seals was tested considering both how the seals learn to produce calls as well as gain information from perceived sounds. In chapter 4, grey seals were shown to be capable of vocal usage learning by producing specific calls in specific contexts. In chapter 5, grey seals displayed control over the structure of their calls as they could imitate novel sound models, demonstrating vocal production learning. Lastly, in chapter 6 grey seals gained information from perceived sound signals and used them to their benefit to locate food, demonstrating acoustic comprehension learning. Adult grey seals have complex call repertoires, and in this thesis I have shown how their calls develop with physiological growth and experience, as well as how they can gain information from perceived sound signals.

During the 1960´s and 1970´s the number of grey seals in the Baltic Sea was decreasing rapidly, mostly due to hunting and toxic substances like DDT and PCB. When hunting became less intense and toxic substances decreased in the environment the grey seal population started to increase. Today grey seals are found common in the Baltic Sea and have started to become a big treat and a problem to the fishing industry. The grey seal destroys and enters fishing traps and consumes large quantities of the fish that have been caught. The knowledge of the grey seal, like abundance and food preferences, is today limited. It is also important to define the position of the grey seal in the ecosystem in the Baltic Sea and to be able to predict changes that could occur if the population would rapidly decrease or increase. This project was therefore started in an attempt to increase the knowledge about the grey seals food preferences. The study was made in tree separate parts. Part one contained analyses of prey remains from stomachs and digestive tract from fourteen individuals put down in two geographically separate areas. The collected material from the seal digestive tract was cleaned and otoliths (hearing stones from fish), scales and back vertebra from fish eaten by the grey seal were sorted out. With the help of hard parts collected from the intestines the food preferences of the seals could be estimated. Eight different species of prey was found. The species were herring (Clupea harengus membras), sprat (Sprattus sprattus), common whitefish (Coregonus spp), perch (Perca fluviatilis), salmon (Salmo salar), trout (Salmo trutta) and roach (Rutilus rutilus). In two of the digestive tracts several individuals of the isopod Saduria entomon were found. No earlier studies describe the isopod as a food source for the grey seals in the Baltic Sea. The findings are therefore unique information. The results showed that during summer the main part of the grey seal diet in the gulf of Sundsvall and in the surrounding coastal area of Hårte was herring and sprat. No significant difference in food preferences was shown between the investigated seals from the gulf of Sundsvall and seals from the surrounding coastal area of Hårte. The second part was made to investigate if it was possible to build a floating platform that would work as a resting place for the grey seal. The surface of the platform was covered by a layer that keeps the seal scats on the platform so that it could be collected. Otoliths from herring and common whitefish were found on the floating platform. Because the platform could not be under surveillance during the whole study some uncertainties about whether the otoliths found came from grey seal or from resting cormorants or other fish eating birds. However, the otolit size is linearly related to the fish size and this relationship can be used to track the predator. Otoliths from herring taken by grey seals and otoliths found on the platform were significantly bigger than the otoliths originating from the prey of cormorants. The results indicated that the common whitefish size was too big for a full grown cormorant bird to consume. The common whitefish size showed that it probably not had been cormorants that had deposited the otoliths on the platform. The platform method was concluded promising but it needs to be modified in order to work more effective in the future. In the third part scats were collected from the area of Österåsen to increase the amount of information about the grey seals food preferences. The knowledge of the grey seals diet in the Baltic Sea is today limited and few similar study’s have earlier been made. The collected scats and otoliths in this project are therefore unique. 2008:Bi 2

<p>During the 1960´s and 1970´s the number of grey seals in the Baltic Sea was decreasing rapidly, mostly due to hunting and toxic substances like DDT and PCB. When hunting became less intense and toxic substances decreased in the environment the grey seal population started to increase. Today grey seals are found common in the Baltic Sea and have started to become a big treat and a problem to the fishing industry. The grey seal destroys and enters fishing traps and consumes large quantities of the fish that have been caught.</p><p>The knowledge of the grey seal, like abundance and food preferences, is today limited. It is also important to define the position of the grey seal in the ecosystem in the Baltic Sea and to be able to predict changes that could occur if the population would rapidly decrease or increase. This project was therefore started in an attempt to increase the knowledge about the grey seals food preferences. The study was made in tree separate parts. Part one contained analyses of prey remains from stomachs and digestive tract from fourteen individuals put down in two geographically separate areas. The collected material from the seal digestive tract was cleaned and otoliths (hearing stones from fish), scales and back vertebra from fish eaten by the grey seal were sorted out. With the help of hard parts collected from the intestines the food preferences of the seals could be estimated. Eight different species of prey was found. The species were herring (Clupea harengus membras), sprat (Sprattus sprattus), common whitefish (Coregonus spp), perch (Perca fluviatilis), salmon (Salmo salar), trout (Salmo trutta) and roach (Rutilus rutilus). In two of the digestive tracts several individuals of the isopod Saduria entomon were found. No earlier studies describe the isopod as a food source for the grey seals in the Baltic Sea. The findings are therefore unique information. The results showed that during summer the main part of the grey seal diet in the gulf of Sundsvall and in the surrounding coastal area of Hårte was herring and sprat. No significant difference in food preferences was shown between the investigated seals from the gulf of Sundsvall and seals from the surrounding coastal area of Hårte.</p><p>The second part was made to investigate if it was possible to build a floating platform that would work as a resting place for the grey seal. The surface of the platform was covered by a layer that keeps the seal scats on the platform so that it could be collected. Otoliths from herring and common whitefish were found on the floating platform. Because the platform could not be under surveillance during the whole study some uncertainties about whether the otoliths found came from grey seal or from resting cormorants or other fish eating birds. However, the otolit size is linearly related to the fish size and this relationship can be used to track the predator. Otoliths from herring taken by grey seals and otoliths found on the platform were significantly bigger than the otoliths originating from the prey of cormorants. The results indicated that the common whitefish size was too big for a full grown cormorant bird to consume. The common whitefish size showed that it probably not had been cormorants that had deposited the otoliths on the platform. The platform method was concluded promising but it needs to be modified in order to work more effective in the future.</p><p>In the third part scats were collected from the area of Österåsen to increase the amount of information about the grey seals food preferences.</p><p>The knowledge of the grey seals diet in the Baltic Sea is today limited and few similar study’s have earlier been made. The collected scats and otoliths in this project are therefore unique.</p><p>2008:Bi 2</p>

Abstract The aims of this thesis included identifying ways to mitigate the economic losses of the commercial fishery in the Gulf of Bothnia resulting from fishery regulation enforced to protect wild Atlantic salmon stocks, the recent decline in survival of hatchery-reared salmon, and the dramatically increased seal-induced catch and gear damage. In addition, these solutions should facilitate safeguarding the recently-recovered wild salmon stocks and seal populations. Other goals included adding to and updating basic knowledge on the effects of increased seal populations, migrating salmon stocks, the coastal trap-net fishery, and their interactions. Seal-induced damage to the commercial fishery was found to be a significant problem throughout the Gulf of Bothnia. Catch and gear damage varied considerably among regions, fishing periods, target species and trap-net models. The regional patterns in seal-induced damage depended on the number of seals in the region and the type of gear, which is strongly associated with the netting materials. Strong and thick materials are more resistant to the attacks of hunting seals. Finer materials and larger mesh sizes that entangle fish are most prone to seal damage. Besides careful selection of the netting material it is also possible to markedly reduce damage by appropriate gear modifications. Three modified trap-net models showed promising results in terms of seal protection, with the pontoon trap being the most successful design. The total size of the spawning salmon population in the Gulf of Bothnia was c. 230 000 in the first two years of the 2000s. The proportions of wild salmon and hatchery-reared salmon, however, appeared to markedly change between years. The survival rate of cultured smolts seems to be considerably lower than that of wild smolts. Large variation in the returning migration patterns and run timing of salmon was found between sea age groups, stock components, and among and within regions. Run timing estimates revealed that the temporal regulation effectively safeguards the wild salmon, but, at the same time, a substantial proportion of the reared salmon escape the coastal fisheries in different regions. The likelihood of survival of wild salmon captured with trap-nets and then released was high, and the cumulative mortality even after several capture-and-release events was estimated to below. Trap-net capture and release did not lead to considerable changes in the normal migration behaviour of Atlantic salmon. This result suggests the potential for a selective harvesting strategy; a system where exploitable (e.g. hatchery-reared fish) and safeguarded (e.g. wild vulnerable stocks) fractions of a fish population complex could be separated in a mixed-stock fishery. However, before introducing a practical and successful selective trap-net fishery, several preconditions should be fulfilled.

Neonatal mortality in grey seals on the Isle of May breeding colony and in a rehabilitation centre were investigated by detailed systematic post-mortem examinations (n=59), on-site bacteriology and advanced molecular diagnostic techniques for specific pathogens. Causes of death on the breeding colony included starvation (30%), omphalitis-peritonitis (26%), septicaemia (22%), stillbirth (10%) and trauma (4%) and in the rehabilitation centre starvation (44%) and septicaemia (22%). Detailed key gross and histopathological findings and pathogens are described and include the first report of Listeria monocytogenes in any marine mammal. Phocid herpes virus 1 nucleic acids were detected in nasal swabs of 58% live, free-ranging grey seal pups (n=90) and 28% yearlings (n=19), suggesting recrudescence in the latter. Previously undetected in Scotland, phocid herpes virus 2 nucleic acids were identified only in yearlings (15%); sealpox was detected in a single live stranded grey seal pup and phocine distemper virus was not detected. Given their unique characteristics and potential for acting as sentinels of coastal marine health several pathogens of putative anthropogenic origin were investigated: Toxoplasma gondii, Neospora caninum, Salmonella spp. and Campylobacter spp. Toxoplasma gondii DNA was detected in 6% dead free-ranging grey seal pups (n=50) but N. caninum was not found. Salmonella (20%) and Campylobacter (50%) were isolated from rectal swabs of live and dead grey seal pups and Campylobacter was significantly associated with moderate to severe colitis implying pathogenicity. These findings imply a land-sea-land transfer of T. gondii and early exposure of pups to this parasite. Extensive genetic fingerprinting suggested an exchange of Salmonella between grey seal, cattle and human populations and that the Campylobacter isolates may share the same origin as human clinical isolates. This work provides a solid base line study of diseases present in grey seal pups and demonstrates that they are useful indicators of coastal marine microbial contamination.

<p>The effects of environmental contaminants in humans and animals are of great concern. Some contaminants are endocrine disruptors that may interfere with the endogenous hormonal signalling and disturb, for example, reproductive organs and functions.</p><p>Primary uterine myometrial cells originating from women and Baltic grey seals were exposed to some polychlorinated biphenyls (PCBs) and their metabolites. Even though human and Baltic grey seal myometrial cells responded differently to the tested PCBs, the results indicate that PCBs can influence myometrial cell proliferation <i>in vitro</i>.</p><p>The prevalence of uterine leiomyomas was investigated among 257 Baltic grey seals. Leiomyomas were only present in females older than 22 years, at a prevalence of 65%. Proliferation in leiomyoma cells was detected in individuals lacking ovarian proliferation support, suggesting the presence of an exogenous stimulant. By taking into account temporal alterations in the contaminant burden of the seals, PCB exposure was found to be associated with leiomyoma prevalence. In conclusion, PCB exposure may be related to uterine leiomyoma development and proliferation in Baltic grey seals <i>in vivo</i>.</p><p>Human endometrial endothelial cells (HEECs) were exposed to some endocrine disruptors, and the effects of the endocrine disruptors on cell proliferation and viability were studied. All evaluated endocrine disruptors decreased HEEC proliferation and most also decreased HEEC viability. Further studies revealed that the reduction in HEEC proliferation after exposure to o,p’-DDT was associated with differential expression of mRNA involved in proliferation, defence response, and lipid and cholesterol metabolism compared to untreated HEEC. </p><p>In conclusion, these studies suggest that endocrine disruptors affect cultured cells from the female reproductive tract of humans and grey seals, and may have deleterious effects on proliferation, viability, and genes involved in defence response, and lipid or cholesterol metabolism.</p>

The effects of environmental contaminants in humans and animals are of great concern. Some contaminants are endocrine disruptors that may interfere with the endogenous hormonal signalling and disturb, for example, reproductive organs and functions. Primary uterine myometrial cells originating from women and Baltic grey seals were exposed to some polychlorinated biphenyls (PCBs) and their metabolites. Even though human and Baltic grey seal myometrial cells responded differently to the tested PCBs, the results indicate that PCBs can influence myometrial cell proliferation in vitro. The prevalence of uterine leiomyomas was investigated among 257 Baltic grey seals. Leiomyomas were only present in females older than 22 years, at a prevalence of 65%. Proliferation in leiomyoma cells was detected in individuals lacking ovarian proliferation support, suggesting the presence of an exogenous stimulant. By taking into account temporal alterations in the contaminant burden of the seals, PCB exposure was found to be associated with leiomyoma prevalence. In conclusion, PCB exposure may be related to uterine leiomyoma development and proliferation in Baltic grey seals in vivo. Human endometrial endothelial cells (HEECs) were exposed to some endocrine disruptors, and the effects of the endocrine disruptors on cell proliferation and viability were studied. All evaluated endocrine disruptors decreased HEEC proliferation and most also decreased HEEC viability. Further studies revealed that the reduction in HEEC proliferation after exposure to o,p’-DDT was associated with differential expression of mRNA involved in proliferation, defence response, and lipid and cholesterol metabolism compared to untreated HEEC. In conclusion, these studies suggest that endocrine disruptors affect cultured cells from the female reproductive tract of humans and grey seals, and may have deleterious effects on proliferation, viability, and genes involved in defence response, and lipid or cholesterol metabolism.

The Optimal Foraging Theory, rooted in the processual archaeology, uses a measuring methodology where the foraging strategy that gives the highest payoff measured as the highest ratio of energy gain per time unit is analysed (Mac Arthur &amp; Pianca 1966, Emlen 1966). The theory is a branch of evolutionary ecology why much attention is paid to the interdependence of humans and preys and environmental conditions caused by climatologically and geographical changes or by overexploitation or other changes caused by humans. The analysis of Early Mesolithic pioneers onGotland, who settle in a transforming landscape, leaves indications of a Maglemose culture origin, probably from flooded original settlements in the south/southwest Baltic basin. The pioneers have to adapt to a seal-hunting economy dominated by grey seal which give the best cost-benefit outcome as big terrestrial mammals are missing and only mountain hare is available. The diet is narrow and there is a great risk for deficiency diseases as well as for acquiring hypervitaminosis and osteoporosis caused by excess of seal food. There is a hiatus c. 5000-4500 BC in the archaeological records on Gotland and the south-western Baltic region, and the master thesis hypothesises that Littorina Transgression I with a severe cold dip called the “8.2 ka BP cold event” has a delayed, but such a severe impact also on fauna and flora on Gotland, that the ecological system is destroyed. The possibility for humans to survive in a sustainable society is questionable. The extremely cold winters during this c. 400 years cold event, with glaciers moving southwards, delayed the blooming season, diminished the harvest and changed both flora and fauna. When the ecological niche for the grey seal is destroyed with flooded beaches close to the pioneers, human overexploitation is reinforced. With a diminishing population of mountain hare, which eventually gets extinct at the end of the Mesolithic, there are no alternatives but some birds and fish, hard to catch. Probably the pioneers abandon Gotland or move to a higher level on Gotland but no records are yet found why the period is called a hiatus. Extinction is the worst scenario or survival in such a small number that a sustainable society is lost. If so, new population groups repopulatedGotlandafter the Littorina transgressions. The origin is still unknown of the Pitted-ware and Funnelbeaker cultures that are populating Gotland after the transgressions. This master thesis can not confirm an affluent life style but rather a suffering starving society flooded by Littorina transgressions and struggling with the severe cold, caused by the “8.2 ka cold event” that makes the environmental conditions even worse. The subsistence economy is successively destroyed which probably causes the hiatus in archaeological records. The Littorina Transgression I with the “8.2 cold event” and the lack of terrestrial big animals are bottle necks. / Optimal Foraging Theory, med sina rötter i den processuella arkeologin, använder en metodik utgående från mätningar där insamlingsstrategin som ger den högsta avkastningen per tidsenhet analyseras (Mac Arthur &amp; Pianca 1966, Emlen 1966). Teorin är en undergrupp inom den evolutionära ekologin och därför ägnas stor tid åt att uppmärksamma det ömsesidiga beroendet och påverkan som sker i miljön p.g.a. klimatologiska och geologiska orsaker men också p.g.a. mänsklig påverkan som exempelvis överförbrukning.  Analysen av tidigmesolitiska pionjärbosättare på Gotland, som möter ett landskap i förvandling, lämnar spår efter sig som tyder på ett ursprung i Maglemosekulturen i södra/sydvästra Östersjöregionen. De tvingas bli adapterade till en säljägarekonomi dominerad av gråsäl som ger det bästa energiutbytet, eftersom stora landdäggdjur saknas och endast bergshare finns tillgänglig. Dietvalet är smalt och det föreligger stor risk för både bristsjukdomar och A-vitaminförgiftning och osteoporos p.g.a. överkonsumtion av sälprodukter. Det finns ett uppehåll i de arkeologiska fynden c. 5000-4500 BC på Gotland liksom i södra Östersjöområdet. Magisteruppsatsens hypotes är att den kalla perioden med temperatursänkning som kallas ”8.2 ka BP cold event” under Littorinatransgression I har en fördröjd men så kraftigt övergripande effekt, på både djur- och växtliv på Gotland, att den förstör det ekologiska systemet och därmed möjligheten för människor att överleva i ett hållbart samhälle. De mycket hårda vintrarna under de c. 400 årens ”cold event” medför att glaciärerna dras sig söderut, blomningssäsongen fördröjs, skörden minskar och både fauna och flora förändras. När den ekologiska nischen för gråsälen förstörs av översvämmade stränder nära bosättarna förstärks överexploateringen, och då det inte finns någon alternativ föda utom en minskande harstam, svårfångade fåglar och fiskar, blir situationen fatal för de tidigmesolitiska bosättarna. Troligtvis flyttar de till andra platser inom Östersjönätverket eller till en högre nivå på Gotland, men fynd saknas hittills varför detta tomrum benämns ”hiatus”. Det värsta scenariot är att bosättarna dör ut eller överlever i ett så litet antal att det hållbara samhället går under.Om så är fallet återbefokas Gotland av gropkeramisk kultur och trattbägarkultur i anslutning till Littorinatrasgressionernas slut. Denna magisteruppsats kan inte konfirmera en livsstil i överflöd, utan snarare ett lidande svältande samhälle som översvämmas av Littorinatransgressioner med miljömässiga förhållanden som förvärras av den allvarliga kylan orsakad av ”8.2 ka cold event”.  Försörjningsmöjligheterna förstörs succesivt och befolkningen försvinner vilket troligen orsakar ett hiatus i de arkeologiska fynden. Littorina Transgression I med ”8.2 ka cold event” och bristen på stora landdjur är stora flaskhalsar.

Recent research using stable isotope analysis has shown a dependence on migrating or breeding populations of vertebrates as vectors for the transfer of marine-derived nutrients within coastal ecosystems. Sable Island, Nova Scotia, Canada supports numerous species of plants, a variety of seabird colonies (including common [Sterna hirundo] and Arctic [Sterna paradisaea] terns), the world’s largest grey seal (Halichoerus grypus) breeding colony, and a self-sustaining population of wild (feral) horses (Equus ferus caballus). I hypothesize that nitrogen cycling within this island ecosystem is highly influenced by the input of nutrients from seals and seabirds (‘biogenic vectors’), affecting primary production and potentially stabilizing higher trophic levels (i.e., horses). To examine this relation I developed a spatially-explicit isoscape for Sable Island through stable isotope analysis of nitrogen (δ15N) in samples (n = 282) of marram grass (Ammophila breviligulata). I incorporated significant variables (i.e., distance to vector colony and distance to shoreline, r2 = 0.41) into the final parsimonious interpolation model using universal co-kriging techniques. The greatest 15N enrichment occurred within the tips and along the perimeter of the island, coinciding with greater densities of grey seals, while the lowest values occurred within the centre of the island. I then identified individual contributions of seal-, tern- and horse-mediated transfer of marine-derived nutrients inland. Marram grass exhibited higher δ15N within seal (μ = 7.5‰) and tern (μ = 5‰) colonies, while horses and biogeochemical processes (i.e., volatilization, ammonification, etc.) most likely contributed to the homogeneity within the centre of the island (μ = 3.6‰). Due to the higher densities, wider distribution, and greater 15N enrichment of marram tissues, grey seals appear to be the most important vector species while seabirds have a more localized effect. The greater availability of N within vector colonies supplemented the local vegetation community, contributing to greater vegetation cover within colony boundaries. This relation had secondary effects on the horse population, which showed correspondingly higher horse δ15N values within the tips of the island (δ15N + 1.6‰) due to consumption of 15N enriched forage. I conclude that biogenic vector species promote nutrient transfer by establishing nutrient gateways which indirectly cause cascading effects throughout the food web.