Environmental change and human history in the Jabal Harun area, Jordan

The research is related to the Finnish Jabal Harun Project (FJHP), which is part of the research unit directed by Professor Jaakko Frösén. The project consists of two interrelated parts: the excavation of a Byzantine monastery/pilgrimage centre on Jabal Harun, and a multiperiod archaeological survey of the surrounding landscape. It is generally held that the Near Eastern landscape has been modified by millennia of human habitation and activity. Past climatic changes and human activities could be expected to have significantly changed also the landscape of the Jabal Harun area. Therefore it was considered that a study of erosion in the Jabal Harun area could shed light on the environmental and human history of the area. It was hoped that it would be possible to connect the results of the sedimentological studies either to wider climatic changes in the Near East, or to archaeologically observable periods of human activity and land use. As evidence of some archaeological periods is completely missing from the Jabal Harun area, it was also of interest whether catastrophic erosion or unfavourable environmental change, caused either by natural forces or by human agency, could explain the gaps in the archaeological record. Changes in climate and/or land-use were expected to be reflected in the sedimentary record. The field research, carried out as part of the FJHP survey fieldwork, included the mapping of wadi terraces and cleaning of sediment profiles which were recorded and sampled for laboratory analyses of facies and lithology. To obtain a chronology for the sedimentation and erosion phases also OSL (optically stimulated luminescence) dating samples were collected. The results were compared to the record of the Near Eastern palaeoclimate, and to data from geoarchaeological studies in central and southern Jordan. The picture of the environmental development was then compared to the human history in the area, based on archaeological evidence from the FJHP survey and the published archaeological research in the Petra region, and the question of the relationship between human activity and environmental change was critically discussed. Using the palaeoclimatic data and the results from geoarchaeological studies it was possible to outline the environmental development in the Jabal Harun area from the Pleistocene to the present.It is appears that there was a phase of accumulation of sediment before the Middle Palaeolithic period, possibly related to tectonic movement. This phase was later followed by erosion, tentatively suggested to have taken place during the Upper Palaeolithic. A period of wadi aggradation probably occurred during the Late Glacial and continued until the end of the Pleistocene, followed by significant channel degradation, attributed to increased rainfall during the Early Holocene. It seems that during the later Holocene channel incision has been dominant in the Jabal Harûn area although there have been also small-scale channel aggradation phases, two of which were OSL-dated to around 4000-3000 BP and 2400-2000 BP. As there is no evidence of tectonic movements in the Jabal Harun area after the early Pleistocene, it is suggested that climate change and human activity have been the major causes of environmental change in the area. At a brief glance it seems that many of the changes in the settlement and land use in the Jabal Harun area can be explained by climatic and environmental conditions. However, the responses of human societies to environmental change are dependent on many factors. Therefore an evaluation of the significance of environmental, cultural, socio-economic and political factors is needed to decide whether certain phenomena are environmentally induced. Comparison with the wider Petra region is also needed to judge whether the phenomena are characteristic of the Jabal Harun area only, or can they be connected to social, political and economic development over a wider area.

J. L. Runebergs samlade skrifter i textkritisk belysning

The Collected Works of J. L. Runeberg from the Viewpoint of Textual Scholarship The theoretical framework of this dissertation builds on textual scholarship. The dissertation explores the history of Runeberg’s publications and his relations with his publishers, from his debut and the first editions, through the editions of collected works published during the course of his life, to the later commercial editions, including the critical edition, published in 1933–2005 by the Svenska Vitterhetssamfundet (The Swedish Society for Belles Lettres) and The Svenska litteratursällskapet i Finland (The Society of Swedish Literature in Finland). The various editions of Runeberg’s collected works are situated in their respective critical traditions, from the 19th century German Ausgabe letzter Hand, to the influence of Anglo-American bibliography on Swedish textual criticism in the late 20th century. By making use of primary material previously not used for research purposes, the author is able to present a new view on Runebergian publishing history, including Runeberg’s fees and his relations with the censor authorities. There are indications that his Finnish publishers could not bear the cost of his sizable fees, that were in proportion neither with the book market in Finland nor with the numbers of copies sold. Apart from a certain body of editions the primary material is comprised of correspondences, publishing contracts, printing house invoices, as well as censor authority records. One of the conclusions drawn is that the early and detailed biography, Biografiska anteckningar om Johan Ludvig Runeberg (Biographical Notes on …) by J. E. Strömborg is not reliable in matters concerning publishing history, and that this work has been used far too uncritically. The history of the critical edition gets a chapter of its own, based on primary material in Swedish and Finnish archives. Finally, the author analyses the critical choices, made primarily in the critical edition, and uses examples from the commercial editions to study the editors’ interventions over time, from the 1850s to the 1920s. The changes to the text are usually small and subtle, but cumulative – and in some cases, crucial for the interpretation of the work. One objective of textual scholarship should be to examine the publishing history of a single work or of an author’s œuvre, and another to pay attention both to changes in a work as such and to the shifts of meaning they might entail.

The effects of human induced disturbances on the sediment geochemistry of Baltic Sea embayments with a focus on eutrophication history

Historical sediment nutrient concentrations and heavy-metal distributions were studied in five embayments in the Gulf of Finland and an adjacent lake. The main objective of the study was to examine the response of these water bodies to temporal changes in human activities. Sediment cores were collected from the sites and dated using 210Pb and 137Cs. The cores were analyzed for total carbon (TC), total nitrogen (TN), total phosphorus (TP), organic phosphorus (OP), inorganic phosphorus (IP), biogenic silica (BSi), loss on ignition (LOI), grain size, Cu, Zn, Al, Fe, Mn, K, Ca, Mg and Na. Principal component analysis (PCA) was used to summarize the trends in the geochemical variables and to compare trends between the different sites. The links between the catchment land use and sediment geochemical data were studied using a multivariate technique of redundancy analysis (RDA). Human activities produce marked geochemical variations in coastal sediments. These variations and signals are often challenging to interpret due to various sedimentological and post-depositional factors affecting the sediment profiles. In general, the sites studied here show significant upcore increases in sedimentation rates, TP and TN concentrations. Also Cu, which is considered to be a good indicator of anthropogenic influence, showed clear increases from 1850 towards the top part of the cores. Based on the RDA-analysis, in the least disturbed embayments with high forest cover, the sediments are dominated by lithogenic indicators Fe, K, Al and Mg. In embayments close to urban settlement, the sediments have high Cu concentrations and a high sediment Fe/Mn ratio. This study suggests that sediment accumulation rates vary significantly from site to site and that the overall sedimentation can be linked to the geomorphology and basin bathymetry, which appear to be the major factors governing sedimentation rates; i.e. a high sediment accumulation rate is not characteristic either to urban or to rural sites. The geochemical trends are strongly site specific and depend on the local geochemical background, basin characteristics and anthropogenic metal and nutrient loading. Of the studied geochemical indicators, OP shows the least monotonic trends in all studied sites. When compared to other available data, OP seems to be the most reliable geochemical indicator describing the trophic development of the study sites, whereas Cu and Zn appear to be good indicators for anthropogenic influence. As sedimentation environments, estuarine and marine sites are more complex than lacustrine basins with multiple sources of sediment input and more energetic conditions in the former. The crucial differences between lacustrine and estuarine/coastal sedimentation environments are mostly related to Fe. P sedimentation is largely governed by Fe redox-reactions in estuarine environments. In freshwaters, presence of Fe is clearly linked to the sedimentation of other lithogenic metals, and therefore P sedimentation and preservation has a more direct linkage to organic matter sedimentation.

Sangen ialo Rucous : Schwenckfeldiläisten rukouskirja Mikael Agricolan lähteenä

Sangen ialo Rucous. The prayer book of the Schwenckfeldiens as a source for Michael Agricola The significance of the prayer book published by the Finnish reformer Michael Agricola in the year of 1544 has not been comprehended enough among the prayer literature of the Reformation century. Especially in foreign research literature one of the era s most extensive and versatile prayer books has been practically disregarded. According to the prayer book, Agricola appears to be a traditionalist, who derives most of his source material from medieval and old church prayer books. The low number of the prayers from Evangelical prayer books is noteworthy. The study in hand examines Agricola s theology expressed in his translation work of the prayer book of the mystic spiritualistic Schwenckfeldien movement. The prayer book of the Schwenckdeldiens diverged from Lutheranism was called Bekantnus der sünden and it was composed around the year of 1526. Agricola is the only prayer book collector who has regarded it as necessary to add all the 45 prayers of the Schwenckfeldien prayer book and its introduction to his book. In the prayers containing the Schwenckfeldien communion theology and Christology Agricola has not changed the content of the text, apparently because he was not aware of the problems involved in the Schwenckfeldien theology. On a few occasions Agricola added points of view concerning the church, priesthood, preaching the Word and sacraments to his prayers, which the Schwenckfeldiens despised. From the ten additions four Agricola created himself, the rest he borrowed from Wolfgang Capito s prayer book. As a source Agricola used Capito s Latin prayer book together with the German text from Bekantnus. When looking at the studied material, Agricola does not turn out to be a creative translator. Even though he had a model for a less restricted translation in Wolfgang Capito s prayer book, he sticks to his habit of translating word by word. Because not even a good example has had a liberating effect on his translation principles, Agricola cannot be considered a theologically orientated writer. The translation of the Schwenckfeldien prayers Agricola starts with Capito s prayer book. Very soon he takes the Bekantnus alongside Capito s text and abandons the use of Capito s prayer book in the middle of the translation process. Comparing Agricola s translation with the two texts in different languages has made it possible to create the disposition theory described above. On the basis of the disposition theory it can be concluded that Agricola first worked out a precise plan for his prayer book. Then he translated prayers theme by theme using multiple sources at the same time. Later Agricola fixed the disposition, which does not seem to have a direct paragon. From the prayers of the Schwenckfeldien prayer book Agricola translated 26 texts using the Bekantnus as an only base, from Capito s prayer book he translated four texts and 17 texts he translated using the two basic texts simultaneously. In the previous studies, words and texts added by Agricola have been examined as one problem unit. In this study the additions have been placed into three different categories: additions consequential on tautological parataxis, specifying additions and additions significant to the content. Due to Agricola s meticulous translation techniques, there are so few additions. Agricola does not show his own creativity even in the additions significant to the content but uses there some complete sentences or word fragments from the other prayers he has translated. In the translations of the prayers there are some unique appearing words and the analysis of the translation work shed a new light on the background of their first literary appearance in Finnish. Agricola s linguistic abilities turned out to be great. In those prayers where Agricola uses both the German and Latin basic texts at the same time, the translation process is a very intensive twine made on the basis of the two sources.

Ihanteena vapaaehtoisuus : Diakoniatyö Porvoon hiippakunnan seurakunnissa, erityisesti Iitin ja Tampereen rovastikunnissa vuosina 1897-1923

The Ideal of Volunteerism. An institutional approach to social welfare work in the parishes of the Diocese of Porvoo especially in the deaneries of Iitti and Tampere, Finland, in the years 1897-1923 Social welfare work (also known as diakonia) has achieved a high status in the Evangelical Lutheran Church of Finland. Since 1944, provisions of the Finnish Church Act have obliged each parish to employ at least one deacon or deaconess. This study sets out to examine the background and development of social welfare work in the Evangelical Lutheran Church of Finland from the 1890s to the 1920s, by which time social welfare work had become an established practice in the Church. The study investigates the development of social welfare work on the level of parishes. The main source material was collected from sixteen parishes in the Diocese of Porvoo especially in the deaneries of Iitti and Tampere. In the 1890s, two approaches were used in church social work in Finland. The dioceses of Kuopio, Savonlinna and Turku pursued a congregational approach to social work, while the Diocese of Porvoo employed an institutional approach, mainly because of the influence of Bishop Herman Råbergh. This study charts the formation of church social work in Finnish parishes, which took place during a period of tension between the two approaches. The institutional approach to church social work adopted by the Diocese of Porvoo was based on the German system of Asisters= houses@, in which deaconess institutes sent parish sisters to serve congregations. The parish or, in many cases, a separate association dedicated to church social work paid an annual fee to the deaconess institute, which took care of the parish sisters in old age. In the institutional approach, volunteers were recruited to carry out church social work. It was considered as inappropriate to use tax revenue or other public funding for church social work, which was supposed to be based on Christian love for one=s fellow humans and the needy, and for which only voluntary financial contributions were supposed to be used. In the congregational approach, church social work was directly based on the efforts of the parish. The approach relied on the administrative bodies of parishes and the Church, and tax revenue collected by the parishes, as well as other forms of public funding, could be used to carry out the social welfare work. The parishes employed deacons and deaconesses and paid their salaries. The approaches described above were not pursued in their ideal forms; instead, many variations existed. However, in principle, the social welfare work undertaken by the parishes of the Diocese of Porvoo was based on the institutional approach, while the congregational approach was largely employed elsewhere in Finland. Both of the approaches were viable. Parishes began to employ deacons and deaconesses as of the 1890s. The number of parishes which had hired a deacon or deaconess increased particularly in the 1910s, by which time 60% of parishes had employed one. This level was maintained until 1944 when each parish in the Evangelical Lutheran Church of Finland was obliged to employ a deacon or deaconess. Deaconesses usually worked as travelling nurses. The autonomous status of Finland as part of the Russian Empire did not give Finns the right to develop legislation on social affairs and health care. Consequently, the legislation process did not begin until Finland gained its independence in 1917. The social welfare work carried out by parishes and a number of voluntary organisations satisfied the emerging need for medical treatment in Finnish society. Neither the government nor the municipalities had sufficient resources to provide this treatment. Based on the ideal of volunteerism, the institutional social work practiced in the Diocese of Porvoo ran into serious difficulties at the end of the First World War. Because of severe inflation, prices began to rise as of 1915 and tripled in 1917-1918. During the same period, Finnish society went through a deep crisis which escalated into Civil War in spring 1918. This period of economic and social turmoil marked a turning-point which led to a weakening of the status of institutional social work in parishes. Voluntary efforts were no longer sufficient to maintain the practice. In contrast, congregational social work, which was based on public funding, was able to cope with the changes and survived the crisis. The approach to social work adopted by the Diocese of Porvoo turned out to be no more than a brief detour in the history of social work in the Evangelical Lutheran Church of Finland. At the start of the 1920s, the two approaches were integrated into a common vision for establishing church social work as a statutory practice in parishes.

Juutalaisvastaisuus suomalaisissa aikakauslehdissä ja kirjallisuudessa 1918-1944

Anti-Semitism existed in Finland during the whole period covered by this study. The immoral acts associated with Jews in the articles were mostly regarded as universal habits, qualities and/or modes of action, that is, unconnected with any particular Finnish Jew. Researchers have tried to explain anti-Semitism in several ways. The theory of Jews as outsiders has been a popular explanation as well as xenophobia, chimerical anti-Semitism and the socio-economic models. The main sources of this study have been over 400 Finnish periodicals and magazines, literature and text books published between 1918 and 1944. This vast number of magazines includes those of the army and the civil guard, religion, humour and the papers of the Finnish extreme right. One can see a distinct foreign and especially German influence in the subjects and phraseology of Finnish anti-Semitic writings between 1918 and 1944. Several known Finnish anti-Semitic writers had some kind of link with Germany. Some Finnish organisations and societies were openly anti-Semitic during this period. There had been cycles in the activity of anti-Semitic writing in Finland, obvious peaks appearing in 1918 1919, 1929 1931, 1933 1938 and 1942 1944. The reason for the 1918 1919 activity was the civil rights which were granted to the Jews in Finland, and the Russian Bolshevik revolution. The worldwide depression from 1929 to 1932 seem to be the reason for new anti-Semitic writing activity. The rise of National Socialism in Germany and the influence this phenomenon had in Finland was the reason for the peak during 1933 1938. During the continuation war 1942 1944 National Socialist Germany was fighting side-by-side with Finland and their anti-Semitic propaganda found easier access to Finland. Of the 433 magazines, journals and newspapers which were used in this study, 71 or 16.4 per cent had at least one article that can be identified as anti-Semitic; especially the magazines of national socialists and other extreme right parties were making anti-Semitic annotations. There were about 50 people known to have written anti-Semitic articles. At least half of these known writers had studied at the university, including as many as 10 priests. Over and above these, there was an even larger number of people who wrote under a pseudonym. The material used suggested that anti-Semitism was not very popular in Finland between 1918 and 1944. Anti-Semitic articles appeared mostly in the magazines of the extreme right, but their circulation was not very large. A proof of the slight influence of these extreme right anti-Semitic ideas is that, beside the tightening of policy towards Jewish immigrants in 1938 and the handing over of eight of these refugees to Germany in 1942, the official policy of Finland never became anti-Semitic. As was stated before, despite the cycles in the number of writings, there does not appear to have been any noticeable change in public opinion. One must also remember that most Finns had not at that period actually met a Jew. The material used suggests that between 1918 and 1944 the so-called Jewish question was seemingly unimportant for most Finns and their attitude to Jews and Jewishness can be described as neutral.

Itsenäisen Suomen puolesta : Sotilaskomitea 1915-1918

For Independent Finland. The Military Committee 1915–1918 In the course of the First World War, several organizations were founded with the purpose of making Finland independent or, at least, restoring her autonomous status. The Military Committee was the most significant active independence organization in Finland in the First World War, in addition to the activist student movement, i.e., the Jaeger Movement. The Military Committee was an organization founded in 1915 by officers who had attended the Hamina Cadet School, with the goal of creating a national army for a liberation war against the Russian troops. It was believed that the liberation war should succeed only with the help of the German Army. With the situation in society continually tensing up in the autumn 1917, the Military Committee also had to figure on the possibility of a Civil War. The activities of the Military Committee started in the early part of 1915 when they were still small-scale, but they gained significant momentum after the Russian Revolution in March 1917. In January 1918, the Military Committee formed the general staff for the White Army, the Senate’s troops. The independence-related activities of the Hamina cadets in the years of the First World War were more extensive and multifaceted than has been believed heretofore. The work of the Military Committee was divided into preparations for a liberation war in Finland, on one hand, and in Stockholm and Berlin, on the other hand. In Finland, the Military Committee took part in intelligence gathering for Germany and in supporting the recruiting Jaegers, and later in founding the civil guard organization, in solving the law and order authorities issue, and finally in selecting the Commander-in-Chief for the Senate’s troops. The member of the Military Committee, especially Captain Hannes Ignatius of the Cavalry contributed greatly to the drafting of the independence activists’ national action plan in Stockholm in May 1917. This plan preceded the formation of the civil guard organization. The Military Committee’s role in founding the civil guards was initially minor, but in the fall of 1917, the Military Committee started to finance the activities of the civil guards, named several former officers as commanders of the civil guards and finally overtook the entire civil guard movement. In Stockholm and Berlin, the representatives of the Military Committee were in active contact with both the high command of the German Army and with the representatives of the Swedish Army. Colonel Nikolai Mexmontan, who was a representative of the Military Committee, collaborated with Swedish officers and Jaeger officers in Stockholm in coming up with comprehensive and detailed plans for starting the Liberation War. Under Mexmontan’s leadership, there were serious negotiations to enter into a confederation with Germany. Lieutenant Colonel Wilhelm Thesleff, on the other hand, became the commander of the Jaeger Battalion 27. The influence and importance of the Military Committee came to the forefront in independent and conflict-torn Finland. The Military Committee became a Senate committee on the 7th of January 1918, with its chairman, for all practical purposes, as the Commander-in-Chief in an eventual war. Lieutenant General Claes Charpentier was the chairman of the Military Committee from mid-December 1917 onwards, but on the 15th of January 1918 he had to resign in favour of Lieutenant General Gustaf Mannerheim. Soon after that, Mannerheim got an order from the chairman of the Senate P. E. Svinhufvud to organize and assume the leadership of the law and order authorities. The chairman of the Military Committee became the Commander-in-Chief of the Senate troops in January 1918, and the Military Committee became the Commander-in-Chief’s general staff. The Military Committee had turned from a clandestine organization into the first general staff of the independent Finnish Army.

Climate forcing on avian life history

In the 21st century, human-induced global climate change has been highlighted as one of the most serious threats to ecosystems worldwide. According to global climate scenarios, the mean temperature in Finland is expected to increase by 1.8 4.0°C by the end of the century. The regional and seasonal change in temperature has predicted to be spatially and temporally asymmetric, where the High-Arctic and Antarctic areas and winter and spring seasons have been projected to face the highest temperature increase. To understand how species respond to the ongoing climate change, we need to study how climate affects species in different phases of their life cycle. The impact of climate on breeding and migration of eight large-sized bird species was studied in this thesis, taking food availability into account. The findings show that climatic variables have considerable impact on the life-history traits of large-sized birds in northern Europe. The magnitude of climatic effects on migration and breeding was comparable with that of food supply, conventionally regarded as the main factor affecting these life-history traits. Based on the results of this thesis and the current climate scenarios, the following not mutually exclusive responses are possible in the near future. Firstly, asymmetric climate change may result in a mistiming of breeding because mild winters and early spring may lead to earlier breeding, whereas offspring are hatching into colder conditions which elevate mortality. Secondly, climate induced responses can differ between species with different breeding tactics (income vs. capital breeding), so that especially capital breeders can gain advantage on global warming as they can sustain higher energy resources. Thirdly, increasing precipitation has the potential to reduce the breeding success of many species by exposing nestlings to more severe post-hatching conditions and hampering the hunting conditions of parents. Fourthly, decreasing ice cover and earlier ice-break in the Baltic Sea will allow earlier spring migration in waterfowl. In eiders, this can potentially lead to more productive breeding. Fifthly, warming temperatures can favour parents preparing for breeding and increase nestling survival. Lastly, the climate-induced phenological changes in life history events will likely continue. Furthermore, interactions between climate and food resources can be complex and interact with each other. Eiders provide an illustrative example of this complexity, being caught in the crossfire between more benign ice conditions and lower salinity negatively affecting their prime food resource. The general conclusion is that climate is controlling not only the phenology of the species but also their reproductive output, thus affecting the entire population dynamics.

Dispersal and related life history traits in the Glanville fritillary butterfly

Most studies of life history evolution are based on the assumption that species exist at equilibrium and spatially distinct separated populations. In reality, this is rarely the case, as populations are often spatially structured with ephemeral local populations. Therefore, the characteristics of metapopulations should be considered while studying factors affecting life history evolution. Theoretical studies have examined spatial processes shaping the evolution of life history traits to some extent, but there is little empirical data and evidence to investigate model predictions. In my thesis I have tried to bridge the gap between theoretical and empirical studies by using the well-known Glanville fritillary (Melitaea cinxia) metapopulation as a model system. The long-term persistence of classic metapopulations requires sufficient dispersal to establish new local populations to compensate for local extinctions. Previous studies on the Glanville fritillary have shown that females establishing new populations are not a random sample from the metapopulation, but they are in fact more dispersive than females in old populations. Many other life-history traits, such as body size, fecundity and lifespan, may be related to dispersal rate. Therefore, I examined a range of correlated traits for their evolutionary and ecological consequences. I was particularly interested in how the traits vary under natural environmental conditions, hence all studies were conducted in a large (32 x 26 m) outdoor population cage built upon a natural habitat patch. Individuals for the experiments were sampled from newly-established and old populations within a large metapopulation. Results show that females originating from newly-established populations had higher within-habitat patch mobility than females from old populations. I showed that dispersal rate is heritable and that flight activity is related to variation in a gene encoding the glycolytic enzyme phosphoglucose isomerase. Both among-individual and among-population variation in dispersal are correlated with the reproductive performance of females, though I found no evidence for a trade-off between dispersal and fecundity in terms of lifetime egg production or clutch size. Instead, the results suggest that highly dispersive females from newly-established populations have a shorter lifespan than females from old populations, and that dispersive females may pay a cost in terms of reduced lifetime reproductive success due to increased time spent outside habitat patches. In summary, the results of this thesis show that genotype-dependent dispersal rate correlates with other life history traits in the Glanville fritillary, and that the rapid turnover of local populations (extinctions and re-colonisations) is likely to be the mechanism that maintains phenotypic variation in many life history traits at the metapopulation level.

Human genetic variation in the Baltic Sea region: Features of population history and natural selection

In this thesis, the genetic variation of human populations from the Baltic Sea region was studied in order to elucidate population history as well as evolutionary adaptation in this region. The study provided novel understanding of how the complex population level processes of migration, genetic drift, and natural selection have shaped genetic variation in North European populations. Results from genome-wide, mitochondrial DNA and Y-chromosomal analyses suggested that the genetic background of the populations of the Baltic Sea region lies predominantly in Continental Europe, which is consistent with earlier studies and archaeological evidence. The late settlement of Fennoscandia after the Ice Age and the subsequent small population size have led to pronounced genetic drift, especially in Finland and Karelia but also in Sweden, evident especially in genome-wide and Y-chromosomal analyses. Consequently, these populations show striking genetic differentiation, as opposed to much more homogeneous pattern of variation in Central European populations. Additionally, the eastern side of the Baltic Sea was observed to have experienced eastern influence in the genome-wide data as well as in mitochondrial DNA and Y-chromosomal variation – consistent with linguistic connections. However, Slavic influence in the Baltic Sea populations appears minor on genetic level. While the genetic diversity of the Finnish population overall was low, genome-wide and Y-chromosomal results showed pronounced regional differences. The genetic distance between Western and Eastern Finland was larger than for many geographically distant population pairs, and provinces also showed genetic differences. This is probably mainly due to the late settlement of Eastern Finland and local isolation, although differences in ancestral migration waves may contribute to this, too. In contrast, mitochondrial DNA and Y-chromosomal analyses of the contemporary Swedish population revealed a much less pronounced population structure and a fusion of the traces of ancient admixture, genetic drift, and recent immigration. Genome-wide datasets also provide a resource for studying the adaptive evolution of human populations. This study revealed tens of loci with strong signs of recent positive selection in Northern Europe. These results provide interesting targets for future research on evolutionary adaptation, and may be important for understanding the background of disease-causing variants in human populations.

Phylogeography and hybrid swarms: history of brackish water bivalve diversity in North European marginal seas

This study addressed the large-scale molecular zoogeography in two brackish water bivalve molluscs, Macoma balthica and Cerastoderma glaucum, and genetic signatures of the postglacial colonization of Northern Europe by them. The traditional view poses that M. balthica in the Baltic, White and Barents seas (i.e. marginal seas) represent direct postglacial descendants of the adjacent Northeast Atlantic populations, but this has recently been challenged by observations of close genetic affinities between these marginal populations and those of the Northeast Pacific. The primary aim of the thesis was to verify, quantify and characterize the Pacific genetic contribution across North European populations of M. balthica and to resolve the phylogeographic histories of the two bivalve taxa in range-wide studies using information from mitochondrial DNA (mtDNA) and nuclear allozyme polymorphisms. The presence of recent Pacific genetic influence in M. balthica of the Baltic, White and Barents seas, along with an Atlantic element, was confirmed by mtDNA sequence data. On a broader temporal and geographical scale, altogether four independent trans-Arctic invasions of Macoma from the Pacific since the Miocene seem to have been involved in generating the current North Atlantic lineage diversity. The latest trans-Arctic invasion that affected the current Baltic, White and Barents Sea populations probably took place in the early post-glacial. The nuclear genetic compositions of these marginal sea populations are intermediate between those of pure Pacific and Atlantic subspecies. In the marginal sea populations of mixed ancestry (Barents, White and Northern Baltic seas), the Pacific and Atlantic components are now randomly associated in the genomes of individual clams, which indicates both pervasive historical interbreeding between the previously long-isolated lineages (subspecies), and current isolation of these populations from the adjacent pure Atlantic populations. These mixed populations can be characterized as self-supporting hybrid swarms, and they arguably represent the most extensive marine animal hybrid swarms so far documented. Each of the three swarms still has a distinct genetic composition, and the relative Pacific contributions vary from 30 to 90 % in local populations. This diversity highlights the potential of introgressive hybridization to rapidly give rise to new evolutionarily and ecologically significant units in the marine realm. In the south of the Danish straits and in the Southern Baltic Sea, a broad genetic transition zone links the pure North Sea subspecies M. balthica rubra to the inner Baltic hybrid swarm, which has about 60 % of Pacific contribution in its genome. This transition zone has no regular smooth clinal structure, but its populations show strong genotypic disequilibria typical of a hybrid zone maintained by the interplay of selection and gene flow by dispersing pelagic larvae. The structure of the genetic transition is partly in line with features of Baltic water circulation and salinity stratification, with greater penetration of Atlantic genes on the Baltic south coast and in deeper water populations. In all, the scenarios of historical isolation and secondary contact that arise from the phylogeographic studies of both Macoma and Cerastoderma shed light to the more general but enigmatic patterns seen in marine phylogeography, where deep genetic breaks are often seen in species with high dispersal potential.