Norm and difference : Stone Age dietary practice in the Baltic region

Stone Age research on Northern Europe frequently makes gross generalizations about the Mesolithic and Neolithic, although we still lack much basic knowledge on how the people lived. The transition from the Mesolithic to the Neolithic in Europe has been described as a radical shift from an economy dominated by marine resources to one solely dependent on farming. Both the occurrence and the geographical extent of such a drastic shift can be questioned, however. It is therefore important to start out at a more detailed level of evidence in order to present the overall picture, and to account for the variability even in such regional or chronological overviews. Fifteen Stone Age sites were included in this study, ranging chronologically from the Early Mesolithic to the Middle or Late Neolithic, c. 8300–2500 BC, and stretching geographically from the westernmost coast of Sweden to the easternmost part of Latvia within the confines of latitudes 55–59° N. The most prominent sites in terms of the number of human and faunal samples analysed are Zvejnieki, Västerbjers and Skateholm I–II. Human and faunal skeletal remains were subjected to stable carbon and nitrogen isotope analysis to study diet and ecology at the sites. Stable isotope analyses of human remains provide quantitative information on the relative importance of various food sources, an important addition to the qualitative data supplied by certain artefacts and structures or by faunal or botanical remains. A vast number of new radiocarbon dates were also obtained. In conclusion, a rich diversity in Stone Age dietary practice in the Baltic Region was demonstrated. Evidence ranging from the Early Mesolithic to the Late Neolithic show that neither chronology nor location alone can account for this variety, but that there are inevitably cultural factors as well. Food habits are culturally governed, and therefore we cannot automatically assume that people at similar sites will have the same diet. Stable isotope studies are very important here, since they tell us what people actually consumed, not only what was available, or what one single meal contained. We should not be deceived in inferring diet from ritually deposited remains, since things that were mentally important were not always important in daily life. Thus, although a ritual and symbolic norm may emphasize certain food categories, these may in fact contribute very little to the diet. By the progress of analysis of intra-individual variation, new data on life history changes have been produced, revealing mobility patterns, breastfeeding behaviour and certain dietary transitions. The inclusion of faunal data has proved invaluable for understanding the stable isotope ecology of a site, and thereby improve the precision of the interpretations of human stable isotope data. The special case of dogs, though, demonstrates that these animals are not useful for inferring human diet, since, due to the number of roles they possess in human society, dogs could deviate significantly from humans in their diet, and in several cases have been proved to do so. When evaluating radiocarbon data derived from human and animal remains from the Pitted-Ware site of Västerbjers on Gotland, the importance of establishing the stable isotope ecology of the site before making deductions on reservoir effects was further demonstrated. The main aim of this thesis has been to demonstrate the variation and diversity in human practices, challenging the view of a “monolithic” Stone Age. By looking at individuals and not only at populations, the whole range of human behaviour has been accounted for, also revealing discrepancies between norm and practice, which are frequently visible both in the archaeological record and in present-day human behaviour.

Deposit-feeding in benthic macrofauna : Tracer studies from the Baltic Sea

A low content of organic matter, which is largely refractory in nature, is characteristic of most sediments, meaning that aquatic deposit-feeders live on a very poor food source. The food is derived mainly from sedimenting phytodetritus, and in temperate waters like the Baltic Sea, from seasonal phytoplankton blooms. Deposit-feeders are either bulk-feeders, or selective feeders, which preferentially ingest the more organic-rich particles in the sediment, including phytodetritus, microbes and meiofauna. The soft-bottom benthos of the Baltic Sea has low species biodiversity and is dominated by a few macrobenthic species, among which the most numerous are the two deposit-feeding amphipods Monoporeia affinis and Pontoporeia femorata, and the bivalve Macoma balthica. This thesis is based on laboratory experiments on the feeding of these three species, and on the priapulid Halicryptus spinulosus. Feeding by benthic animals is often difficult to observe, but can be effectively studied by the use of tracers. Here we used the radioactive isotope 14C to label food items and to trace the organic matter uptake in the animals, while the stable isotopes 13C and 15N were used to follow feeding on aged organic matter in the sediment. The abundance of M. balthica and the amphipods tends to be negatively correlated, i.e., fewer bivalves are found at sites with dense populations of amphipods, with the known explanation that newly settled M. balthica spat are killed by the amphipods. Whether the postlarvae are just accidentally killed, or also ingested after being killed was tested by labelling the postlarvae with 14C and Rhodamine B. Both tracer techniques gave similar evidence for predation on and ingestion of postlarval bivalves. We calculated that this predation was likely to supply less than one percent of the daily carbon requirement for M. affinis, but might nevertheless be an important factor limiting recruitment of M. balthica. The two amphipods M. affinis and P. femorata are partly vertically segregated in the sediment, but whether they also feed at different depths was unknown. By adding fresh 14C-labelled algae either on the sediment surface or mixed into the sediment, we were able to distinguish surface from subsurface feeding. We found M. affinis and P. femorata to be surface and subsurface deposit-feeders, respectively. Whether the amphipods also feed on old organic matter, was studied by adding fresh 14C-labelled algae on the sediment surface, and using aged, one-year-old 13C- and 15N-labelled sediment as deep sediment. Ingestion of old organic matter, traced by the stable isotopes, differed between the two species, with a higher uptake for P. femorata, suggesting that P. femorata utilises the older, deeper-buried organic matter to a greater extent. Feeding studies with juveniles of both M. affinis and P. femorata had not been done previously. In an experiment with the same procedure and treatments as for the adults, juveniles of both amphipod species were found to have similar feeding strategies. They fed on both fresh and old sediment, with no partitioning of food resources, making them likely to be competitors for the same food resource. Oxygen deficiency has become more wide-spread in the Baltic Sea proper in the last half-century, and upwards of 70 000km2 are now devoid of macrofauna, even though part of that area does not have oxygen concentrations low enough to directly kill the macrofauna. We made week-long experiments on the rate of feeding on 14C-labelled diatoms spread on the sediment surface in different oxygen concentrations for both the amphipod species, M. balthica and H. spinulosus. The amphipods were the most sensitive to oxygen deficiency and showed reduced feeding and lower survival at low oxygen concentrations. M. balthica showed reduced feeding at the lowest oxygen concentration, but no mortality increase. The survival of H. spinulosus was unaffected, but it did not feed, showing that it is not a surface deposit-feeder. We conclude that low oxygen concentrations that are not directly lethal, but reduce food intake, may lead to starvation and death in the longer term.

Factors structuring Fucus communities at open and complex coastlines in the Baltic Sea

This thesis deals with physical factors and biological interactions affecting the distribution of two fucoid species, Fucus vesiculosus and F. serratus, in the Baltic Sea. Studies have been carried out in two quite different environments: an archipelago, and an open rocky coast. The archipelago has an extremely long coastline with a heterogeneous submerged landscape of different substrate types, slopes, water qualities, and degrees of wave exposure. The factors influencing F. vesiculosus distribution, morphology and epiphyte composition were studied in the Stockholm archipelago using field surveys and spatial modelling in Geographic information systems (GIS). A GIS-method to estimate wave exposure was developed and validated by comparing the result to an index based on vertical zonation of lichens. Wave exposure was considered an important factor for predicting the distribution of F. vesiculosus by its ability to clean hard surfaces from silt, and a predictive model was constructed based on the information of wave exposure and slope of the shore. It is suggested that the lower distribution boundary of attached F. vesiculosus is set by sediment in sheltered parts of the archipelago, and by light availability in highly wave exposed parts. The morphology of F. vesiculosus was studied over a wave exposure gradient, and several characters responded in accordance with earlier studies. However, when separating effects of wave exposure from effects of other confounding water property parameters, only thallus width was significantly different. Several water property parameters were shown to be correlated with wave exposure in the Stockholm archipelago, and the mechanism responsible for the effects on F. vesiculosus morphology is discussed. The composition of epiphytes on F. vesiculosus varied over a wave exposure gradient with a positive correlation to Elachista fucicola, and a negative to Chorda filum. At an open coast the physical environment is much less heterogeneous compared to an archipelago. The distributions of F. vesiculosus, F. serratus, turf-forming algae, and the seafloor substrate, were surveyed along the open coasts of Öland and Gotland. Turf-forming algae dominated all hard substrates in the area, and Polysiphonia fucoides was most abundant. At the Gotland coast F. vesiculosus was less abundant than at the Öland coast, and F. serratus occurred only in the southern-most part. Fucus serratus was increasingly more common towards south which was interpreted as an effect mainly of the Baltic salinity gradient, or the variation of salinity that has occurred in the past. The effects of turf-forming algae and sediment on F. serratus recruitment at 7 m depth off the Öland east coast were studied in the field, and by laboratory experiments. Almost no recruits were found in the algal turf outside the F. serratus patches. More fine sediment was found in the turf than in the F. serratus patches, suggesting that the turf accumulates sediment by decreasing resuspension. Both filamentous algae and sediment decreased the attachment ability of F. serratus zygotes and survival of recruits, and sediment had the strongest effect. It is therefore suggested that F. serratus has difficulties recruiting outside its patches, and that these difficulties are enforced by the eutrophication of the Baltic Sea, which has favoured growth of filamentous algae and increased sedimentation. An overall conclusion is that Fucus distribution is affected by large-scale-factors, such as the eutrophication and salinity changes of the Baltic Sea, as well as by small-scale variation in wave exposure, substrate and slope, and by surface competition with neighbouring species.

The Emergence of the Post-Socialist Welfare State - The Case of the Baltic States : Estonia, Latvia and Lithuania

This dissertation takes a step towards providing a better understanding of post-socialist welfare state development from a theoretical as well as an empirical perspective. The overall analytical goal of this thesis has been to critically assess the development of social policies in Estonia, Latvia and Lithuania using them as illustrative examples of post-socialist welfare state development in the light of the theories, approaches and typologies that have been developed to study affluent capitalist democracies. The four studies included in this dissertation aspire to a common aim in a number of specific ways. The first study tries to place the ideal-typical welfare state models of the Baltic States within the well-known welfare state typologies. At the same time, it provides a rich overview of the main social security institutions in the three countries by comparing them with each other and with the previous structures of the Soviet period. It examines the social insurance institutions of the Baltic States (old-age pensions, unemployment insurance, short-term benefits, sickness, maternity and parental insurance and family benefits) with respect to conditions of eligibility, replacement rates, financing and contributions. The findings of this study indicate that the Latvian social security system can generally be labelled as a mix of the basic security and corporatist models. The Estonian social security system can generally also be characterised as a mix of the basic security and corporatist models, even if there are some weak elements of the targeted model in it. It appears that the institutional changes developing in the social security system of Lithuania have led to a combination of the basic security and targeted models of the welfare state. Nevertheless, as the example of the three Baltic States shows, there is diversity in how these countries solve problems within the field of social policy. In studying the social security schemes in detail, some common features were found that could be attributed to all three countries. Therefore, the critical analysis of the main social security institutions of the Baltic States in this study gave strong supporting evidence in favour of identifying the post-socialist regime type that is already gaining acceptance within comparative welfare state research. Study Two compares the system of social maintenance and insurance in the Soviet Union, which was in force in the three Baltic countries before their independence, with the currently existing social security systems. The aim of the essay is to highlight the forces that have influenced the transformation of the social policy from its former highly universal, albeit authoritarian, form, to the less universal, social insurance-based systems of present-day Estonia, Latvia and Lithuania. This study demonstrates that the welfare–economy nexus is not the only important factor in the development of social programs. The results of this analysis revealed that people's attitudes towards distributive justice and the developmental level of civil society also play an important part in shaping social policies. The shift to individualism in people’s mentality and the decline of the labour movement, or, to be more precise, the decline in trade union membership and influence, does nothing to promote the development of social rights in the Baltic countries and hinders the expansion of social policies. The legacy of the past has been another important factor in shaping social programs. It can be concluded that social policy should be studied as if embedded not only in the welfare-economy nexus, but also in the societal, historical and cultural nexus of a given society. Study Three discusses the views of the state elites on family policy within a wider theoretical setting covering family policy and social policy in a broader sense and attempts to expand this analytical framework to include other post-socialist countries. The aim of this essay is to explore the various views of the state elites in the Baltics concerning family policy and, in particular, family benefits as one of the possible explanations for the observed policy differences. The qualitative analyses indicate that the Baltic States differ significantly with regard to the motives behind their family policies. Lithuanian decision-makers seek to reduce poverty among families with children and enhance the parents’ responsibility for bringing up their children. Latvian policy-makers act so as to increase the birth rate and create equal opportunities for children from all families. Estonian policy-makers seek to create equal opportunities for all children and the desire to enhance gender equality is more visible in the case of Estonia in comparison with the other two countries. It is strongly arguable that there is a link between the underlying motives and the kinds of family benefits in a given country. This study, thus, indicates how intimately the attitudes of the state bureaucrats, policy-makers, political elite and researchers shape social policy. It confirms that family policy is a product of the prevailing ideology within a country, while the potential influence of globalisation and Europeanisation is detectable too. The final essay takes into account the opinions of welfare users and examines the performances of the institutionalised family benefits by relying on the recipients’ opinions regarding these benefits. The opinions of the populations as a whole regarding government efforts to help families are compared with those of the welfare users. Various family benefits are evaluated according to the recipients' satisfaction with those benefits as well as the contemporaneous levels of subjective satisfaction with the welfare programs related to the absolute level of expenditure on each program. The findings of this paper indicate that, in Latvia, people experience a lower level of success regarding state-run family insurance institutions, as compared to those in Lithuania and Estonia. This is deemed to be because the cash benefits for families and children in Latvia are, on average, seen as marginally influencing the overall financial situation of the families concerned. In Lithuania and Estonia, the overwhelming majority think that the family benefit systems improve the financial situation of families. It appears that recipients evaluated universal family benefits as less positive than targeted benefits. Some universal benefits negatively influenced the level of general satisfaction with the family benefits system provided in the countries being researched. This study puts forward a discussion about whether universalism is always more legitimate than targeting. In transitional economies, in which resources are highly constrained, some forms of universal benefits could turn out to be very expensive in relative terms, without being seen as useful or legitimate forms of help to families. In sum, by closely examining the different aspects of social policy, this dissertation goes beyond the over-generalisation of Eastern European welfare state development and, instead, takes a more detailed look at what is really going on in these countries through the examples of Lithuania, Latvia and Estonia. In addition, another important contribution made by this study is that it revives ‘western’ theoretical knowledge through ‘eastern’ empirical evidence and provides the opportunity to expand the theoretical framework for post-socialist societies.

Barshalder 1 : A cemetery in Grötlingbo and Fide parishes, Gotland, Sweden, c. AD 1-1100. Excavations and finds 1826-1971

The prehistoric cemetery of Barshalder is located along the main road on the boundary between Grötlingbo and Fide parishes, near the southern end of the island of Gotland in the Baltic Sea. The cemetery was used from c. AD 1-1100. The level of publication in Swedish archaeology of the first millennium AD is low compared to, for instance, the British and German examples. Gotland’s rich Iron Age cemeteries have long been intensively excavated, but few have received monographic treatment. This publication is intended to begin filling this gap and to raise the empirical level of the field. It also aims to make explicit and test the often somewhat intuitively conceived results of much previous research. The analyses deal mainly with the Migration (AD 375–540), Vendel (AD 520–790) and Late Viking (AD 1000–1150) Periods. The following lines of inquiry have been prioritised. 1. Landscape history, i.e. placing the cemetery in a landscape-historical context. (Vol. 1, section 2.2.6) 2. Migration Period typochronology, i.e. the study of change in the grave goods. (Vol. 2, chapter 2) 3. Social roles: gender, age and status. (Vol. 2, chapter 3) 4. Religious identity in the 11th century, i.e. the study of religious indicators in mortuary customs and grave goods, with particular emphasis on the relationship between Scandinavian paganism and Christianity.. (Vol. 2, chapter 4) Barshalder is found to have functioned as a central cemetery for the surrounding area, located on peripheral land far away from contemporary settlement, yet placed on a main road along the coast for maximum visibility and possibly near a harbour. Computer supported correspondence analysis and seriation are used to study the gender attributes among the grave goods and the chronology of the burials. New methodology is developed to distinguish gender-neutral attributes from transgressed gender attributes. Sub-gender grouping due to age and status is explored. An independent modern chronology system with rigorous type definitions is established for the Migration Period of Gotland. Recently published chronology systems for the Vendel and Viking Periods are critically reviewed, tested and modified to produce more solid models. Social stratification is studied through burial wealth with a quantitative method, and the results are tested through juxtaposition with several other data types. The Late Viking Period graves of the late 10th and 11th centuries are studied in relation to the contemporary Christian graves at the churchyards. They are found to be symbolically soft-spoken and unobtrusive, with all pagan attributes kept apart from the body in a space between the feet of the deceased and the end of the over-long inhumation trench. A small number of pagan reactionary graves with more forceful symbolism are however also identified. The distribution of different 11th century cemetery types across the island is used to interpret the period’s confessional geography, the scale of social organisation and the degree of allegiance to western and eastern Christianity. 11th century society on Gotland is found to have been characterised by religious tolerance, by an absence of central organisation and by slow piecemeal Christianisation.

Barshalder 2 : Studies of late Iron Age Gotland

The prehistoric cemetery of Barshalder is located along the main road on the boundary between Grötlingbo and Fide parishes, near the southern end of the island of Gotland in the Baltic Sea. The ceme-tery was used from c. AD 1-1100. The level of publication in Swedish archaeology of the first millennium AD is low compared to, for instance, the British and German examples. Gotland’s rich Iron Age cemeteries have long been intensively excavated, but few have received monographic treatment. This publication is intended to begin filling this gap and to raise the empirical level of the field. It also aims to make explicit and test the often somewhat intuitively conceived re-sults of much previous research. The analyses deal mainly with the Migration (AD 375–540), Vendel (AD 520–790) and Late Viking (AD 1000–1150) Periods. The following lines of inquiry have been prioritised. 1. Landscape history, i.e. placing the cemetery in a landscape-historical context. (Vol. 1, section 2.2.6) 2. Migration Period typochronology, i.e. the study of change in the grave goods. (Vol. 2, chapter 2) 3. Social roles: gender, age and status. (Vol. 2, chapter 3) 4. Religious identity in the 11th century, i.e. the study of religious indicators in mortuary cus-toms and grave goods, with particular emphasis on the relationship between Scandinavian paganism and Christianity. (Vol. 2, chapter 4) Barshalder is found to have functioned as a central cemetery for the surrounding area, located on pe-ripheral land far away from contemporary settle-ment, yet placed on a main road along the coast for maximum visibility and possibly near a harbour. Computer supported correspondence analysis and seriation are used to study the gender attributes among the grave goods and the chronology of the burials. New methodology is developed to distin-guish gender-neutral attributes from transgressed gender attributes. Sub-gender grouping due to age and status is explored. An independent modern chronology system with rigorous type definitions is established for the Migration Period of Gotland. Recently published chronology systems for the Vendel and Viking Periods are critically reviewed, tested and modified to produce more solid models. Social stratification is studied through burial wealth with a quantitative method, and the results are tested through juxtaposition with several other data types. The Late Viking Period graves of the late 10th and 11th centuries are studied in relation to the contemporary Christian graves at the churchyards. They are found to be symbolically soft-spoken and unobtrusive, with all pagan attributes kept apart from the body in a space between the feet of the deceased and the end of the over-long inhumation trench. A small number of pagan reactionary graves with more forceful symbolism are however also identified. The distribution of different 11th cen-tury cemetery types across the island is used to in-terpret the period’s confessional geography, the scale of social organisation and the degree of alle-giance to western and eastern Christianity. 11th century society on Gotland is found to have been characterised by religious tolerance, by an absence of central organisation and by slow piecemeal Christianisation.

Speciation and fractionation of Ca and the REE in fresh and marine waters

This study is concerned with speciation and fractionation of the rare earth elements (REE) and calcium (Ca) in aqueous solutions. The aim is to investigate the chemical states and physical sizes in which these elements can be present. The REE (including neodymium) and Ca have contrasting geochemical behavior in aqueous solutions. Ca is a major dissolved element, while the REE are trace components and highly reactive with aquatic particles. The major interests of the five papers included in this thesis are the following: · Papers I and V deal with the behavior of neodymium (Nd) and its isotopes in the Kalix River and some marine waters. · The diffusive gradients in thin-films (DGT) method is developed for measuring Ca and Mg in Paper II. · Paper III presents a speciation and fractionation study of Ca in the Kalix and Amazonian rivers. · The rare earth elements and their carrier phases are investigated in the Kalix river in Paper IV. For most elements a detailed study of speciation and fractionation can not be performed using only one method. This is due to the overall heterogeneity of the material, considering both size and chemical composition, which is present in aquatic solutions. During this project the aquatic geochemistry of the REE and Ca has been studied using mainly three methods; cross-flow filtration (CFF), field-flow fractionation (FFF) and diffusive gradients in thin-films (DGT). Field work has to a large part been conducted in the Kalix River, in northern Sweden, which is one of the last pristine river systems in Europe. Some field work has also been conducted in the Baltic Sea and the Arctic Ocean. Results from Amazonian rivers are also presented. These are the main conclusions from this work: The DGT technique works equally well for measuring Ca and Mg in natural waters as previously reported for trace metal. A significant colloidal phase for Ca could be detected in the Kalix River and in different Amazonian rivers. This was concluded independently using both CFF and FFF. Variations in REE signatures in the Kalix River suggests two different pathways for the REE during weathering and release form soil profiles and transport in the river. No significant variation in Nd-isotopic composition could be detected in the Kalix River although concentrations varied by a factor of ~10. This suggests that there is one major source for Nd in the river although different pathways for the REE may exist. A study of Nd in the Kalix River, the Baltic Sea and the Arctic Ocean showed that the isotopic compositions in the diffusible fractions were similar to water samples. However, the relative amount of diffusible Nd increased with salinity, probably reflecting the lower concentration of colloidal and particulate material in marine waters.

Dispersal in free-living, marine, benthic nematodes : passive or active processes?

Meiofauna, and especially marine nematodes are common in sediments around the world. Despite very wide ranging distributions in many nematode species, little is presently known about their dispersal mechanisms shaping these patterns. Rafting, and perhaps ballast water transport has been suggested as viable means for nematode long-range transport. On a much smaller scale other processes have been suggested for their dispersal. They generally include some form of passive suspension into the water column and later on a passive, haphazard settling back towards the bottom. Small-scale phenomena in nematode dispersal were studied by conducting a series of studies at Askö field station, Trosa Archipelago, Baltic proper. Studied aspects were one case of macrofaunal influence on nematode dispersal rate, using an amphipod, Monoporeia affinis as disturbing agent, and three different studies on mechanisms related to settling. The experiments were conducted both in laboratory and field settings. The amphipod Monoporeia affinis did not exert any influence on the dispersal rate in the nematodes. The nematode dispersal was only an effect of time, in the aspect that the more time that past, the more nematodes dispersed from their place of origin. The settling experiments revealed that nematodes do have an active component in their settling behaviour, as they were able to exert influence on the spot where they were to settle. They were able to choose settling spot in response to the food quality of the sediment. It also became evident that contrary to common belief, nematodes are able to extend their presence in the water column far beyond the times that would be predicted considering settling velocities and hydrodynamic conditions alone.

An evaluation of the distributions of polychlorinated biphenyls and organic matter in coastal sediments

The objective of this thesis is to improve the understanding of what processes and mechanism affects the distribution of polychlorinated biphenyls (PCBs) and organic carbon in coastal sediments. Because of the strong association of hydrophobic organic contaminants (HOCs) such as PCBs with organic matter in the aquatic environment, these two entities are naturally linked. The coastal environment is the most complex and dynamic part of the ocean when it comes to both cycling of organic matter and HOCs. This environment is characterised by the largest fluxes and most diverse sources of both entities. A wide array of methods was used to study these processes throughout this thesis. In the field sites in the Stockholm archipelago of the Baltic proper, bottom sediments and settling particulate matter were retrieved using sediment coring devices and sediment traps from morphometrically and seismically well-characterized locations. In the laboratory, the samples have been analysed for PCBs, stable carbon isotope ratios, carbon-nitrogen atom ratios as well as standard sediment properties. From the fieldwork in the Stockholm Archipelago and the following laboratory work it was concluded that the inner Stockholm archipelago has a low (≈ 4%) trapping efficiency for freshwater-derived organic carbon. The corollary is a large potential for long-range waterborne transport of OC and OC-associated nutrients and hydrophobic organic pollutants from urban Stockholm to more pristine offshore Baltic Sea ecosystems. Theoretical work has been carried out using Geographical Information Systems (GIS) and statistical methods on a database of 4214 individual sediment samples, each with reported individual PCB congener concentrations. From this work it was concluded that the continental shelf sediments are key global inventories and ultimate sinks of PCBs. Depending on congener, 10-80% of the cumulative historical emissions to the environment are accounted for in continental shelf sediments. Further it was concluded that the many infamous and highly contaminated surface sediments of urban harbours and estuaries of contaminated rivers cannot be of importance as a secondary source to sustain the concentrations observed in remote sediments. Of the global shelf PCB inventory < 1% are in sediments near population centres while ≥ 90% is in remote areas (> 10 km from any dwellings). The remote sub-basin of the North Atlantic Ocean contains approximately half of the global shelf sediment inventory for most of the PCBs studied.

Marine Seaweed Invasions : the Ecology of Introduced Fucus evanescens

Biological invasions are an important issue of global change and an increased understanding of invasion processes is of crucial importance for both conservation managers and international trade. In this thesis, I have studied the invasion of the brown seaweed Fucus evanescens, to investigate the fate and effect of a perennial, habitat-forming seaweed introduced to a coastal ecosystem. A long-term study of the spread of F. evanescens in Öresund (southern Sweden) showed that the species was able to expand its range quickly during the first 20 years after the introduction, but that the expansion has been slow during the subsequent 30 years. Both in Öresund and in Skagerrak, the species is largely restricted to sites where native fucoids are scarce. Laboratory experiments showed that the restricted spread of F. evanescens cannot be explained by the investigated abiotic factors (wave exposure and salinity), although salinity restricts the species from spreading into the Baltic Sea. Neither did I find evidence for that herbivores or epibiota provide biotic resistance to the invader. On the contrary, F. evanescens was less consumed by native herbivores, both compared to the native fucoids and to F. evanescens populations in its native range, and little overgrown by epiphytes. Instead, the restricted spread may be due to competition from native seaweeds, probably by pre-occupation of space, and the establishment has probably been facilitated by disturbance. The studies provided little support for a general enemy release in introduced seaweeds. The low herbivore consumption of F. evanescens in Sweden could not be explained by release from specialist herbivores. Instead, high levels of chemical anti-herbivore defence metabolites (phlorotannins) could explain the pattern of herbivore preference for different fucoids. Likewise, the low epibiotic colonisation of F. evanescens plants could be explained by high resistance to epibiotic survival. This shows that colonisation of invading seaweeds by native herbivores and epibionts depends on properties of the invading species. The large differences between fucoid species in their quality as food and habitat for epibionts and herbivores imply that invasions of such habitat-forming species may have a considerable effect on a number of other species in shallow coastal areas. However, since F. evanescens did not exclude other fucoids in its new range, its effect on the recipient biota is probably small.

The juvenile three-spined stickleback : model organism for the study of estrogenic and androgenic endocrine disruption in laboratory and field

Industrial and domestic sewage effluents have been found to cause reproductive disorders in wild fish, often as a result of the interference of compounds in the effluents with the endocrine system. This thesis describes laboratory-based exposure experiments and a field survey that were conducted with juveniles of the three-spined stickleback, Gasterosteus aculeatus. This small teleost is a common fish in Swedish coastal waters and was chosen as an alternative to non-native test species commonly used in endocrine disruption studies, which allows the comparison of field data with results from laboratory experiments. The aim of this thesis was to elucidate 1) if genetic sex determination and differentiation can be disturbed by natural and synthetic steroid hormones and 2) whether this provides an endpoint for the detection of endocrine disruption, 3) to evaluate the applicability of specific estrogen- and androgen-inducible marker proteins in juvenile three-spined sticklebacks, 4) to investigate whether estrogenic and/or androgenic endocrine disrupting activity can be detected in effluents from Swedish pulp mills and domestic sewage treatment plants and 5) whether such activity can be detected in coastal waters receiving these effluents. Laboratory exposure experiments found juvenile three-spined sticklebacks to be sensitive to water-borne estrogenic and androgenic steroid substances. Intersex – the co-occurrence of ovarian and testicular tissue in gonads – was induced by 17β-estradiol (E2), 17α-ethinylestradiol (EE2), 17α-methyltestosterone (MT) and 5α-dihydrotestosterone (DHT). The first two weeks after hatching was the phase of highest sensitivity. MT was ambivalent by simultaneously eliciting masculinizing and feminizing effects. When applying a DNA-based method for genetic sex identification, it was found that application of MT only during the first two weeks after hatching caused total and apparently irreversible development of testis in genetic females. E2 caused gonad type reversal from male to female. E2 and EE2 induced vitellogenin - the estrogen-responsive yolk precursor protein, while DHT and MT induced spiggin – the androgen-responsive glue protein of the stickleback. None of the effluents from two pulp mills and two domestic sewage treatment plants had any estrogenic or androgenic activity. Juvenile three-spined sticklebacks were collected during four subsequent summers at the Swedish Baltic Sea coast in recipients of effluents from pulp mills and a domestic sewage treatment plant as well as remote reference sites. No sings of endocrine disruption were observed at any site, when studying gonad development or marker proteins, except for a deviation of sex ratios at a reference site. The three-spined stickleback – with focus on the juvenile stage – was found to be a sensitive species suitable for the study of estrogenic and androgenic endocrine disruption.