A sediment record from the western Gotland Basin, northwestern Baltic Proper, covering the last 1200 years, was investigated for past changes in climate and the environment using diatoms as a proxy. The aim is to compare the environmental conditions reconstructed during Medieval times with settings occurring the last century under influence of environmental stressors like eutrophication and climate change. The study core records more marine conditions in the western Gotland Basin surface waters during the Medieval Climate Anomaly (MCA; 950–1250C.E.), with a salinity of at least 8 psu compared to the present 6.5 psu. The higher salinity together with a strong summer-autumn stratification caused by warmer climate resulted in extensive long-lasting diatom blooms of Pseudosolenia calcar-avis, effectively enhancing the vertical export of organic carbon to the sediment and contributing to benthic hypoxia. Accordingly, our data support that a warm and dry climate induced the extensive hypoxic areas in the open Baltic Sea during the MCA. During the Little ice Age (LIA; 1400–1700C.E.), the study core records oxic bottom water conditions, decreasing salinity and less primary production. This was succeeded during the 20th century, about 1940, by environmental changes caused by human-induced eutrophication. Impact of climate change is visible in the diatom composition data starting about 1975C.E. and becoming more pronounced 2000C.E., visible as an increase of taxa that thrived in stratified waters during autumn blooms typically due to climate warming.

This study aims to investigate and disentangle the impact of land use and climate variability on the Baltic Sea coastal zone from a millennial perspective. To assess the environmental status of the coastal zone we make use of siliceous microfossils (mainly diatoms), stable nitrogen and carbon isotopes, organic carbon accumulation rates, and lithological changes analyzed in a sediment core collected in Gamlebyviken, Swedish east coast, dated to cover the last 3000 years. Changes in land use and vegetation cover are modelled using pollen stratigraphical data to obtain the percentage coverage of coniferous woodland (Pinus and Picea), deciduous woodland, wetland (Cyperaceae), grassland (including Juniperus) and cropland (cereals) while changes in climatic conditions are assessed through well-documented climatic periods that have occurred in the Baltic Sea region. The reconstructed regional vegetation cover shows that already 3000 years ago, humans used the landscape for both animal husbandry (grasslands) and farming (cropland), but the impact on the Baltic coastal waters was minor. The diatom accumulation rates were quite high (similar to 3100-2600 cal yr BP) containing taxa indicative of high nutrient conditions/upwelling, and stable carbon isotopes show that the carbon was produced in the basin but did not result in elevated organic carbon accumulation rates. A gradual change to less marine conditions in Gamlebyviken from about 2500 to 1400 cal yr BP can be attributed to the ongoing land uplift which resulted in a more enclosed embayment with only a narrow inlet area today. The Medieval Climate Anomaly (1000-700 cal yr BP/950-1250 CE) is a time where extensive eutrophication is registered in the open Baltic Sea, but afforestation is recorded between 1000 and 500 cal yr BP and attributed to the expansion of spruce favored by land-use reorganization with a transition from a one-course rotation system to the three-course rotation system fully established in southern Sweden in the 13th century, and only minor environmental change is recorded in the coastal zone. The Little Ice Age is documented in our data between 400 and 250 cal yr BP/1550-1700 CE as a decrease in regional cropland (cereals) cover, possibly indicating years of poor crop harvest, and changes in the Baltic coastal zone are evidenced as low carbon and diatom accumulation rates, increase in benthic diatom taxa (low turbidity), and high abundance in diatom taxa associated with sea ice indicating a cold climate. The most significant changes occurred from about 100 cal yr BP/1850 CE up to present, with a maximum regional cover of grassland and cropland (ca. 35%) at the expense of deciduous woodland, and major changes indicative of a highly eutrophic environment recorded in the coastal zone. Organic carbon accumulation rates peaked in 1968 CE at approximately 134 g C m(2) yr(-1) before subsequently declining to present-day values of 53 g C m(2) yr(-1), mirroring a similar trend observed in diatom accumulation rates. The high organic carbon accumulation rate shows that deep unvegetated accumulation bottoms in the coastal Baltic Sea serve as carbon sinks and are worth exploring for their potential in mitigating climate change. Variation partitioning shows that 26% of the variance in the diatom assemblages is associated with land use changes. The variables grassland, cropland, and stable nitrogen isotopes are accordingly strong predictors of environmental change in the Baltic coastal zone as reflected by the diatom assemblages.

This study aims to investigate and disentangle the impact of land use and climate variability on the Baltic Sea coastal zone from a millennial perspective. To assess the environmental status of the coastal zone we make use of siliceous microfossils (mainly diatoms), stable nitrogen and carbon isotopes, organic carbon accumulation rates, and lithological changes analyzed in a sediment core collected in Gamlebyviken, Swedish east coast, dated to cover the last 3000 years. Changes in land use and vegetation cover are modelled using pollen stratigraphical data to obtain the percentage coverage of coniferous woodland (Pinus and Picea), deciduous woodland, wetland (Cyperaceae), grassland (including Juniperus) and cropland (cereals) while changes in climatic conditions are assessed through well-documented climatic periods that have occurred in the Baltic Sea region. The reconstructed regional vegetation cover shows that already 3000 years ago, humans used the landscape for both animal husbandry (grasslands) and farming (cropland), but the impact on the Baltic coastal waters was minor. The diatom accumulation rates were quite high (∼3100–2600 cal yr BP) containing taxa indicative of high nutrient conditions/upwelling, and stable carbon isotopes show that the carbon was produced in the basin but did not result in elevated organic carbon accumulation rates. A gradual change to less marine conditions in Gamlebyviken from about 2500 to 1400 cal yr BP can be attributed to the ongoing land uplift which resulted in a more enclosed embayment with only a narrow inlet area today. The Medieval Climate Anomaly (1000–700 cal yr BP/950–1250 CE) is a time where extensive eutrophication is registered in the open Baltic Sea, but afforestation is recorded between 1000 and 500 cal yr BP and attributed to the expansion of spruce favored by land-use reorganization with a transition from a one-course rotation system to the three-course rotation system fully established in southern Sweden in the 13th century, and only minor environmental change is recorded in the coastal zone. The Little Ice Age is documented in our data between 400 and 250 cal yr BP/1550–1700 CE as a decrease in regional cropland (cereals) cover, possibly indicating years of poor crop harvest, and changes in the Baltic coastal zone are evidenced as low carbon and diatom accumulation rates, increase in benthic diatom taxa (low turbidity), and high abundance in diatom taxa associated with sea ice indicating a cold climate. The most significant changes occurred from about 100 cal yr BP/1850 CE up to present, with a maximum regional cover of grassland and cropland (ca. 35%) at the expense of deciduous woodland, and major changes indicative of a highly eutrophic environment recorded in the coastal zone. Organic carbon accumulation rates peaked in 1968 CE at approximately 134 g C m2 yr−1 before subsequently declining to present-day values of 53 g C m2 yr−1, mirroring a similar trend observed in diatom accumulation rates. The high organic carbon accumulation rate shows that deep unvegetated accumulation bottoms in the coastal Baltic Sea serve as carbon sinks and are worth exploring for their potential in mitigating climate change. Variation partitioning shows that 26% of the variance in the diatom assemblages is associated with land use changes. The variables grassland, cropland, and stable nitrogen isotopes are accordingly strong predictors of environmental change in the Baltic coastal zone as reflected by the diatom assemblages.

We have reviewed information on early-, late- and limit-value decomposition stages for litter of Norway spruce (Picea abies) and Scots pine (Pinus silvestris). This synthesis covers c 16 studies/papers made along a climatic gradient; range in mean annual temperature (MAT) from −1 to +7 °C and mean annual precipitation (MAP) from 425 to 1070 mm. Scots pine has an early stage dominated by carbohydrate decomposition and a late stage dominated by decomposition of lignin; Norway spruce has just one stage dominated by lignin decomposition. We used data for annual mass loss to identify rate-regulating factors in both stages; climate data, namely, MAT and MAP, as well as substrate properties, namely, nitrogen (N), acid unhydrolyzable residue (AUR), manganese (Mn). Early-stage decomposition for Scots pine litter was dominated positively by MAT; the late stage was dominated negatively by MAT, N, and AUR, changing with decomposition stage; there was no effect of Mn. Norway spruce litter had no early stage; decomposition in the lignin-dominated stage was mainly negative to MAP, a negative relationship to AUR and non-significant relationships to N and MAT. Mn had a positive relationship. Limit values for decomposition, namely, the accumulated mass loss at which decomposition is calculated to be zero, were related positively to Mn and AUR for Scots pine litter and negatively to AUR for Norway spruce litter. With different sets of rate-regulating factors as well as different compounds/elements related to the limit values, the decomposition patterns or pathways are different.

The aim of this paper is to show different patterns for decomposition of the main mass of needle litter from two boreal and temperate coniferous tree species, both leading to a stabilized fraction of litter. To this purpose we have reviewed information on decomposition patterns in the lignin-dominated (late) stages of two local foliar litter types, namely those of Scots pine (Pinus silvestris) and Norway spruce (Picea abies) from two climatic gradients of equal extension. We have also reviewed factors determining the limit values for both species.

Long-term decomposition studies were used to calculate annual mass loss in the lignin-dominated decomposition stage and relate these to mean annual temperature (MAT), mean annual precipitation (MAP) and concentrations of nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg), manganese (Mn) and acid unhydrolyzable residue (gravimetric lignin, AUR).

There was no effect of MAT on decomposition of either needle litter type. MAP had a rate-dampening effect on decomposition of Norway spruce litter. There was a rate-stimulating effect of Mn for Norway spruce litter but not for that of Scots pine. In spite of the strong negative effect of AUR and N on decomposition of Scots pine litter there was none at all for that of Norway spruce.

Limit values for decomposition were related to the litters’ initial concentrations of N, Mn and AUR and differed between litter types for locally collected, natural litter and for that from experimental litter, the latter having higher N and lower Mn concentrations than the natural litter.

We conclude that the two litter types have clear differences as regards rate- regulating factors for decomposition in the late lignin-dominated stage as well as for the stable fraction and suggest two different pathways for their decomposition. This is the first time that different pathways have been suggested for decomposing litter.

We synthesized available data for magnesium (Mg) dynamics in newly shed and decomposing foliar litter of mainly pine (Pinus) species, Norway spruce (Picea abies), and birch (Betula) species. Using original, measured data from 40 stands organized in climatic gradients we intended to determine patterns of Mg concentration and net release vs accumulated mass loss of the litter. This synthesis is likely the first synthesis of Mg dynamics in decomposing litter.

In paired stands, litter of both Norway spruce and lodgepole pine (Pinus contorta) had higher Mg concentrations than Scots pine (Pinus silvestris), with concentrations in Norway spruce litter even twice as high.

In decomposing litter, Mg concentrations followed a quadratic (X2-X) function vs accumulated mass loss and consequently had minima, different for Norway spruce and Scots pine litter. Out of 68 decomposition studies 53 gave minimum concentration. The Mg minimum concentration during decomposition was positively related to initial Mg concentration for Scots pine and Scots pine plus lodgepole pine but not for Norway spruce. The increase in concentration suggests that after the minimum Mg was temporarily limiting.

For Norway spruce litter there was a relationship between minimum concentration of Mg and the limit value. There was no such relationship for Scots pine and not for the combined pine data.

Magnesium net release started directly after the incubation and was linear to accumulated mass loss of litter, giving a slope coefficient (release rate) for each study. The net release rate was linear to initial Mg concentration and all studies combined gave a negative linear relationship.

Rapid evolutionary change is increasingly being recognized as commonplace, but the evolutionary consequences for species and ecosystems under human-induced selection regimes have not been explored in detail, although many species occur in such environments. In a common garden experiment and with amplified fragment length polymorphism markers, we examined whether genetic differentiation has taken place between spatially intermixed populations of creeping thistles Cirsium arvense (Asteraceae) collected from a natural habitat (maritime shores), a semi-natural habitat (road verges) and arable fields under two management regimes: conventional and organic farming. Populations of C. arvense have altered genetically and locally adapted their growth patterns with changed land use. Although plants from different habitats showed similar total biomass production, shoot and root production was higher for maritime populations, suggesting selection for increased competitive ability. Competitive ability then declined in the order semi-natural, conventional farms and organic farms. Thistles in arable fields may be more selected for tolerance against disturbances from herbicides and mechanical weed control. In addition, early shoot sprouting and genetic analysis showed differentiation between plants originating from conventional farms and farms that were converted to organic 9–30 years ago, suggesting some adaptation to altered crop cultivation practices

Rapid evolutionary change is increasingly being recognized as commonplace, but the evolutionary consequences for species and ecosystems under human-induced selection regimes have not been explored in detail, although many species occur in such environments. In a common garden experiment and with amplified fragment length polymorphism markers, we examined whether genetic differentiation has taken place between spatially intermixed populations of creeping thistles Cirsium arvense (Asteraceae) collected from a natural habitat (maritime shores), a semi-natural habitat (road verges) and arable fields under two management regimes: conventional and organic farming. Populations of C. arvense have altered genetically and locally adapted their growth patterns with changed land use. Although plants from different habitats showed similar total biomass production, shoot and root production was higher for maritime populations, suggesting selection for increased competitive ability. Competitive ability then declined in the order semi-natural, conventional farms and organic farms. Thistles in arable fields may be more selected for tolerance against disturbances from herbicides and mechanical weed control. In addition, early shoot sprouting and genetic analysis showed differentiation between plants originating from conventional farms and farms that were converted to organic 9–30 years ago, suggesting some adaptation to altered crop cultivation practices

Native wild boar (Sus scrofa) populations are expanding across Europe. This is cause for concern in some areas where overabundant populations impact natural ecosystems and adjacent agronomic systems. To better manage the potential for impacts, managers require more information about how the species may affect other organisms. For example, information regarding the effect of wild boar on soil fungi for management application is lacking. Soil fungi play a fundamental role in ecosystems, driving essential ecological functions; acting as mycorrhizal symbionts, sustaining plant nutrition and providing defense; as saprotrophs, regulating the organic matter decomposition; or as plant pathogens, regulating plant fitness and survival. During autumn (Sep–Nov) 2018, we investigated the effects of wild boar (presence/absence and rooting intensity) on the abundance (number of individuals) of fungal sporocarps and their functional guilds (symbiotic, saprotrophic and pathogenic). We selected eleven forested sites (400–500 × 150–200 m) in central Sweden; six with and five without the presence of wild boar. Within each forest, we selected one transect (200 m long), and five plots (2 × 2 m each) for sites without wild boar, and ten plots for sites with boars (five within and five outside wild boar disturbances), to determine the relationship between the intensity of rooting and the abundance of sporocarps for three fungal guilds. We found that the presence of wild boar and rooting intensity were associated with the abundance of sporocarps. Interestingly, this relationship varied depending on the fungal guild analyzed, where wild boar rooting had a positive correlation with saprophytic sporocarps and a negative correlation with symbiotic sporocarps. Pathogenic fungi, in turn, were more abundant in undisturbed plots (no rooting) but located in areas with the presence of wild boar. Our results indicate that wild boar activities can potentially regulate the abundance of fungal sporocarps, with different impacts on fungal guilds. Therefore, wild boar can affect many essential ecosystem functions driven by soil fungi in boreal forests, such as positive effects on energy rotation and in creating mineral availability to plants, which could lead to increased diversity of plants in boreal forests.

Tobacco smoking is a risk factor for numerous disorders, including cancers affecting organs outside the respiratory tract. Epidemiological data suggest that smoking is a greater risk factor for these cancers in males compared to females. This observation, together with the fact that males have a higher incidence of and mortality from most non-sex-specific cancers, remains unexplained. Loss of chromosome Y (LOY) in blood cells is associated with increased risk of nonhematological tumors. We demonstrate here that smoking is associated with LOY in blood cells in three independent cohorts [TwinGene: odds ratio (OR) = 4.3, 95% CI = 2.8-6.7; ULSAM: OR = 2.4, 95% CI = 1.6-3.6; and PIVUS: OR = 3.5, 95% CI = 1.4-8.4] encompassing a total of 6014 men. The data also suggest that smoking has a transient and dose-dependent mutagenic effect on LOY status. The finding that smoking induces LOY thus links a preventable risk factor with the most common acquired human mutation.

Tobacco smoking is a risk factor for numerous disorders, including cancers affecting organs outside the respiratory tract. Epidemiological data suggest that smoking is a greater risk factor for these cancers in males compared to females. This observation, together with the fact that males have a higher incidence of and mortality from most non-sex-specific cancers, remains unexplained. Loss of chromosome Y (LOY) in blood cells is associated with increased risk of nonhematological tumors. We demonstrate here that smoking is associated with LOY in blood cells in three independent cohorts [TwinGene: odds ratio (OR) = 4.3, 95% CI = 2.8-6.7; ULSAM: OR = 2.4, 95% CI = 1.6-3.6; and PIVUS: OR = 3.5, 95% CI = 1.4-8.4] encompassing a total of 6014 men. The data also suggest that smoking has a transient and dose-dependent mutagenic effect on LOY status. The finding that smoking induces LOY thus links a preventable risk factor with the most common acquired human mutation.

Tick-borne encephalitis virus (TBEV) is a flavivirus with major impact on global health. The geographical TBEV distribution is expanding, thus making it pivotal to further characterize the natural virus populations. In this study, we completed the earlier partial sequencing of a TBEV pulled out of a pool of RNA extracted from 115 ticks collected on Torö in the Stockholm archipelago. The total RNA was sufficient for all sequencing of a TBEV genome (Torö-2003), without conventional enrichment procedures such as cell culturing or suckling mice amplification. To our knowledge, this is the first time that the genome of TBEV has been sequenced directly from an arthropod reservoir. The Torö-2003 sequence has been characterized and compared with other TBE viruses. In silico analyses of secondary RNA structures formed by the two untranslated regions revealed a temperature-sensitive structural shift between a closed replicative form and an open AUG accessible form, analogous to a recently described bacterial thermoswitch. Additionally, novel phylogenetic conserved structures were identified in the variable part of the 3'-untranslated region, and their sequence and structure similarity when compared with earlier identified structures suggests an enhancing function on virus replication and translation. We propose that the thermo-switch mechanism may explain the low TBEV prevalence often observed in environmentally sampled ticks. Finally, we were able to detect variations that help in the understanding of virus adaptations to varied environmental temperatures and mammalian hosts through a comparative approach that compares RNA folding dynamics between strains with different mammalian cell passage histories.

Tick-borne encephalitis virus (TBEV) is a flavivirus with major impact on global health. The geographical TBEV distribution is expanding, thus making it pivotal to further characterize the natural virus populations. In this study, we completed the earlier partial sequencing of a TBEV pulled out of a pool of RNA extracted from 115 ticks collected on Torö in the Stockholm archipelago. The total RNA was sufficient for all sequencing of a TBEV genome (Torö-2003), without conventional enrichment procedures such as cell culturing or suckling mice amplification. To our knowledge, this is the first time that the genome of TBEV has been sequenced directly from an arthropod reservoir. The Torö-2003 sequence has been characterized and compared with other TBE viruses. In silico analyses of secondary RNA structures formed by the two untranslated regions revealed a temperature-sensitive structural shift between a closed replicative form and an open AUG accessible form, analogous to a recently described bacterial thermoswitch. Additionally, novel phylogenetic conserved structures were identified in the variable part of the 3'-untranslated region, and their sequence and structure similarity when compared with earlier identified structures suggests an enhancing function on virus replication and translation. We propose that the thermo-switch mechanism may explain the low TBEV prevalence often observed in environmentally sampled ticks. Finally, we were able to detect variations that help in the understanding of virus adaptations to varied environmental temperatures and mammalian hosts through a comparative approach that compares RNA folding dynamics between strains with different mammalian cell passage histories.

Habitats of Swedish conservation interests are in general small and fragmented following the extensive and intensive forest management history. This study covering 71 000 ha of boreal Sweden investigates how history influences present-day distribution and composition of forests identified as high conservation value habitats and how they are protected. We also investigated if the habitat criteria used to describe reservations differed between reservation types and if habitat criteria were associated with the size of Woodland Key Habitats. The results show strong effects from historical ownership and historical forest type on the probability of an area being set aside as formally protected or as voluntary protected Woodland Key Habitats. We also found that both formal reservations and Woodland Key Habitats primarily cover coniferous forest in the age interval 70-110 years but not the presumably most valuable oldest coniferous category >110 or deciduous forests, which are as common in reservations as in other areas. Old deciduous forests (>110 years) are significantly more rare in formal reservations compared to the forest matrix. When viewed in a context of fragmentation and edge effects the results underline the importance of evaluating reserved areas and Woodland Key Habitats in a wider temporal and larger spatial perspective to optimize conservation management efforts. Maximal representation and biodiversity can be better achieved if new reservations are chosen to represent different ownership and forest history, and if they are selected in a landscape context related to present reservations and the present surrounding production forest.

Habitats of Swedish conservation interests are in general small and fragmented following the extensive and intensive forest management history. This study covering 71 000 ha of boreal Sweden investigates how history influences present-day distribution and composition of forests identified as high conservation value habitats and how they are protected. We also investigated if the habitat criteria used to describe reservations differed between reservation types and if habitat criteria were associated with the size of Woodland Key Habitats. The results show strong effects from historical ownership and historical forest type on the probability of an area being set aside as formally protected or as voluntary protected Woodland Key Habitats. We also found that both formal reservations and Woodland Key Habitats primarily cover coniferous forest in the age interval 70-110 years but not the presumably most valuable oldest coniferous category >110 or deciduous forests, which are as common in reservations as in other areas. Old deciduous forests (>110 years) are significantly more rare in formal reservations compared to the forest matrix. When viewed in a context of fragmentation and edge effects the results underline the importance of evaluating reserved areas and Woodland Key Habitats in a wider temporal and larger spatial perspective to optimize conservation management efforts. Maximal representation and biodiversity can be better achieved if new reservations are chosen to represent different ownership and forest history, and if they are selected in a landscape context related to present reservations and the present surrounding production forest.

In this study we reconstruct forest composition during the 1860s for a 71000 ha area in southern boreal Sweden. The aim is to show how historical ownership and associated anthropogenic disturbances act as a source of heterogeneity in the present-day distribution and composition of coniferous and deciduous forest within the commercial production forest. We use older (>110 years) and mature (70-110 years) forest as response variables in generalized linear models with a binominal error distribution. The explanatory variables include size of zone, historical type of ownership zone (village, company, and farm), amount of forest, and forest type. We focus in particular on investigating effects from interacting explanatory variables. The significant statistical associations in the study indicate that patterns of deciduous and coniferous older patches differ, and that deciduous patches differ in relation to age interval. The oldest deciduous patches, for example, are today more likely on areas that had deciduous cover also in the past and stood on forestland managed by farmers, but less likely on the same habitat managed by companies. We show that there are strong effects on present forest composition from historical ownership and forest composition. We argue that by including local data on past ownership combined with knowledge on use patterns management could be better adapted to local landscape dynamics compared to the application of overly generalized patterns or models of boreal dynamics that excludes interactions with management.

In this study we reconstruct forest composition during the 1860s for a 71000 ha area in southern boreal Sweden. The aim is to show how historical ownership and associated anthropogenic disturbances act as a source of heterogeneity in the present-day distribution and composition of coniferous and deciduous forest within the commercial production forest. We use older (>110 years) and mature (70-110 years) forest as response variables in generalized linear models with a binominal error distribution. The explanatory variables include size of zone, historical type of ownership zone (village, company, and farm), amount of forest, and forest type. We focus in particular on investigating effects from interacting explanatory variables. The significant statistical associations in the study indicate that patterns of deciduous and coniferous older patches differ, and that deciduous patches differ in relation to age interval. The oldest deciduous patches, for example, are today more likely on areas that had deciduous cover also in the past and stood on forestland managed by farmers, but less likely on the same habitat managed by companies. We show that there are strong effects on present forest composition from historical ownership and forest composition. We argue that by including local data on past ownership combined with knowledge on use patterns management could be better adapted to local landscape dynamics compared to the application of overly generalized patterns or models of boreal dynamics that excludes interactions with management.

We combine historical maps and satellite derived data to reconstruct the development of a Swedish boreal landscape over the past 300 years. The aim is to understand legacies from past use patterns in present-day forest composition and consequences for conservation objectives from a landscape perspective. We analyze landscape development in cross-tabulation matrixes, building change trajectories. These trajectories are tested in linear models to explain the distribution of present-day landscape composition of coniferous, mixed, and deciduous forests >110 years. Of 49 tested change trajectories, 11 showed a significant association. Associations for mixed and coniferous forests were similar and linked to characteristics such as forest continuity, which characterized the studied landscape. Deciduous older forests did not show any association to forest continuity but were more likely to occur on areas that specifically shifted from forests with grazing in the 1720s to open impediment (likely indicating low tree coverage) in the 1850s. There were large shifts and spatial redistribution in ownerships over time. Use patterns and legacies varied between small- and large-scale ownership categories as well as within small-scale categories. The legacies found in the study indicate a complex origin of heterogeneous landscape elements such as older deciduous forests. Additionally, the origin of the legacies indicates a potential need to diversify conservation management based on the influence of past use patterns. Despite large inconsistencies in historical and contemporary data we argue that this type of analysis could be used to further understand the distribution of landscape elements important for conservation objectives.

We combine historical maps and satellite derived data to reconstruct the development of a Swedish boreal landscape over the past 300 years. The aim is to understand legacies from past use patterns in present-day forest composition and consequences for conservation objectives from a landscape perspective. We analyze landscape development in cross-tabulation matrixes, building change trajectories. These trajectories are tested in linear models to explain the distribution of present-day landscape composition of coniferous, mixed, and deciduous forests >110 years. Of 49 tested change trajectories, 11 showed a significant association. Associations for mixed and coniferous forests were similar and linked to characteristics such as forest continuity, which characterized the studied landscape. Deciduous older forests did not show any association to forest continuity but were more likely to occur on areas that specifically shifted from forests with grazing in the 1720s to open impediment (likely indicating low tree coverage) in the 1850s. There were large shifts and spatial redistribution in ownerships over time. Use patterns and legacies varied between small- and large-scale ownership categories as well as within small-scale categories. The legacies found in the study indicate a complex origin of heterogeneous landscape elements such as older deciduous forests. Additionally, the origin of the legacies indicates a potential need to diversify conservation management based on the influence of past use patterns. Despite large inconsistencies in historical and contemporary data we argue that this type of analysis could be used to further understand the distribution of landscape elements important for conservation objectives.

We studied genetic effects of the colonisation process during primary succession by analysing allozyme variation at a PGI locus in differently aged populations of Moehringia trinervia, which is a selfing annual with low dispersal ability. The populations studied come from islands and shores created in the 1880s by a drop in the water table of a Swedish lake and from old parts of a large island and of the mainland. The population age is known from five vegetation analyses over a century. We have also analysed the genetic composition of M. trinervia derived from seeds in the soil. Mainland populations had a higher genetic diversity than island populations that were little differentiated and differed genetically from the mainland populations. There was no temporal trend in the distribution of genetic variation on the new islands. The presence of alleles in the extant populations was associated with the proportion of that allele in the seed bank, indicating a main recruitment from the seed bank and not by repeated immigrations. We suggest that some of the new islands were colonised by a few early founders from the mainland. Later colonisation has occurred between adjacent islands, which preserves the founder effect and could explain the uniform, low genetic variation in the island populations

We studied genetic effects of the colonisation process during primary succession by analysing allozyme variation at a PGI locus in differently aged populations of Moehringia trinervia, which is a selfing annual with low dispersal ability. The populations studied come from islands and shores created in the 1880s by a drop in the water table of a Swedish lake and from old parts of a large island and of the mainland. The population age is known from five vegetation analyses over a century. We have also analysed the genetic composition of M. trinervia derived from seeds in the soil. Mainland populations had a higher genetic diversity than island populations that were little differentiated and differed genetically from the mainland populations. There was no temporal trend in the distribution of genetic variation on the new islands. The presence of alleles in the extant populations was associated with the proportion of that allele in the seed bank, indicating a main recruitment from the seed bank and not by repeated immigrations. We suggest that some of the new islands were colonised by a few early founders from the mainland. Later colonisation has occurred between adjacent islands, which preserves the founder effect and could explain the uniform, low genetic variation in the island populations

Sphagnum (peatmoss) dominates huge areas of the Northern Hemisphere and acts as a significant carbon sink on a global scale, yet little is known about the genetic structure of Sphagnum populations. We investigated genetic structure within a population of the common peatmoss Sphagnum fuscum, to assess local patterns of genetic diversity and the spatial extent of clones. One hundred seventeen shoots were sampled from five transects in Fuglmyra, central Norway, and sequenced for three anonymous DNA regions. Five neighbourhood patches were marked along each transect, and from each patch, five stems were sampled for molecular analyses. Seventeen haplotypes could be distinguished and two major groups of haplotypes differed by 12 mutational steps. The two major haplotype groups differed significantly in microhabitat association along the distance to groundwater table and the pH gradients, indicating microhabitat differentiation. The haplotypes within these groups were all genetically similar, differing by one or two mutations. The most common haplotype occurred in four transects separated by 250-m distance. Most of the molecular variation in the population was found among transects, and within patches. Large dominating clones within each transect resulted in low variation explained by the among-patch-within-transect component of spatial structure. Mutation appears to account for a larger proportion of the population variation than recombination. Within the population, vegetative growth and asexual reproduction from gametophyte fragments dominate as the main reproductive mode

Sphagnum (peatmoss) dominates huge areas of the Northern Hemisphere and acts as a significant carbon sink on a global scale, yet little is known about the genetic structure of Sphagnum populations. We investigated genetic structure within a population of the common peatmoss Sphagnum fuscum, to assess local patterns of genetic diversity and the spatial extent of clones. One hundred seventeen shoots were sampled from five transects in Fuglmyra, central Norway, and sequenced for three anonymous DNA regions. Five neighbourhood patches were marked along each transect, and from each patch, five stems were sampled for molecular analyses. Seventeen haplotypes could be distinguished and two major groups of haplotypes differed by 12 mutational steps. The two major haplotype groups differed significantly in microhabitat association along the distance to groundwater table and the pH gradients, indicating microhabitat differentiation. The haplotypes within these groups were all genetically similar, differing by one or two mutations. The most common haplotype occurred in four transects separated by 250-m distance. Most of the molecular variation in the population was found among transects, and within patches. Large dominating clones within each transect resulted in low variation explained by the among-patch-within-transect component of spatial structure. Mutation appears to account for a larger proportion of the population variation than recombination. Within the population, vegetative growth and asexual reproduction from gametophyte fragments dominate as the main reproductive mode

Using fast-evolving microsatellites, more slowly evolving ITS markers and performing habitat analyses, we demonstrated a drastic genetic divergence and significant habitat differentiation between early- and late-flowering variants of plants morphologically belonging to Gymnadenia conopsea ssp conopsea. The two phenological variants can either be found in separate or in mixed populations. Information from microsatellite markers and ITS sequences indicated the occurrence of an early historical split between the two flowering-time variants, a split that has been maintained until the present time even within sympatric populations. Early-flowering variants were also far more genetically diverse, had more alleles per microsatellite locus and a wider habitat amplitude than late-flowering variants. As a comparison, we included G. odoratissima in the sequencing study. We found G. odoratissima to be most closely related to the early-flowering type. This indicates a more ancient divergence event between the two flowering-time variants within G. conopsea ssp conopsea than between the two different species G. odoratissima and the early-flowering variant of G. conopsea. Possible explanations to the results arrived at and possible mechanisms maintaining the genetic separation are discussed.

Using fast-evolving microsatellites, more slowly evolving ITS markers and performing habitat analyses, we demonstrated a drastic genetic divergence and significant habitat differentiation between early- and late-flowering variants of plants morphologically belonging to Gymnadenia conopsea ssp conopsea. The two phenological variants can either be found in separate or in mixed populations. Information from microsatellite markers and ITS sequences indicated the occurrence of an early historical split between the two flowering-time variants, a split that has been maintained until the present time even within sympatric populations. Early-flowering variants were also far more genetically diverse, had more alleles per microsatellite locus and a wider habitat amplitude than late-flowering variants. As a comparison, we included G. odoratissima in the sequencing study. We found G. odoratissima to be most closely related to the early-flowering type. This indicates a more ancient divergence event between the two flowering-time variants within G. conopsea ssp conopsea than between the two different species G. odoratissima and the early-flowering variant of G. conopsea. Possible explanations to the results arrived at and possible mechanisms maintaining the genetic separation are discussed.

Sediment cores from three sites along the east-coast of Sweden, north-western Baltic Proper, have been studied with respect to lithologies, geochemistry, and diatom assemblages to trace and date early human impact with emphasis on nutrient discharge. The three sites Braviken, Himmerfjarden, and Adfjarden, have been impacted to various degree during the last millennia by multiple stressors like excessive nutrient discharge and hazardous substances, leading to coastal hypoxia, eutrophication, and pollution. These stressors are mainly caused by drivers in the drainage area as increased human population, changed land use, and point sources as industries and a sewage treatment plant. Even though their detailed history differs, the results show similar general patterns for all three sites. We find no evidence in our data from the coastal zone supporting the hypothesis that the extensive areal distribution of hypoxia in the open Baltic Sea during the Medieval Climate Anomaly was caused by human impact. Timing of the onset of man-made eutrophication, as identified from delta N-15 and changes in diatom composition, differs between the three sites, reflecting the site specific geography and local environmental histories of these areas. The onset of eutrophication dates to similar to 1800 CE in Braviken and Himmerfjarden areas, and to similar to 1900 CE in the less urban area of Adfjarden. We conclude that the recorded environmental changes during the last centuries are unique in a thousand year perspective.

Vicia pisiformis L. is a perennial leguminous plant with a main distribution in broadleaved forest-steppes of eastern Europe. The species is classified as endangered (EN) according to the IUCN red-lists in both Norway and Sweden, due to severe fragmentation, small population sizes and continuing population decline. The populations on the Scandinavian Peninsula constitute the northern limit of the species distribution and are mostly restricted to warm stony slopes with predominantly southern aspects. In this study we used the AFLP method, which is a high-resolution genetic fingerprint method. Samples were collected from 22 Scandinavian populations. The overall genetic structure was analysed in an AMOVA, in a Mantel test and through constrained correspondence analysis (CCA). The ordination scores representing non-geographic genetic divergence were extracted from the CCA and analysed in a linear model using habitat variables and population size as explanatory variables. We found (i) a strong geographic structure, (ii) significant genetic divergence between populations, (iii) that this genetic divergence remained significant even after removing the effect of geography in a partial CCA and (iv) that the remaining non-geographic part of genetic divergence (distance from the ordination centre) was associated with aspect, populations with a northern aspect were more genetically divergent. Aspect explains more variation than population size and is the only variable retained in the minimal adequate model. We suggest that local adaptation has caused this divergence from an expected geographical pattern of genetic variation. This explanation is further supported by the association between aspect and specific AFLP fragments. Many plant populations are relics of a different climate (Aguirre-Planter et al. 2000; Despres et al. 2002; Pico and Riba 2002). In response to long-term climate change, populations can either migrate towards a more favourable climate or adapt to the new conditions (delaVega 1996; Jump et al. 2006). Species with limited dispersal ability are at risk of reaching isolated dead-ends of decreasingly suitable habitat, without any suitable habitat within dispersal distance (Colas et al. 1997). Isolated populations have to use their inherent evolutionary potential and adapt to changes in environmental conditions, or they will go extinct. As population fragments go extinct, those that remain will become increasingly isolated from each other both spatially and also genetically as the level of gene flow declines with increasing distance. Such correlation between genetic dissimilarities and geographic distances, known as isolation by distance (Slatkin 1993; Wright 1943), when found, suggests a history of geographically limited gene flow (Kimura and Weiss 1964). On top of an isolation by distance pattern there might be other genetic structures to be found. Occasional long-distance dispersal events for example may disturb geographic patterns with puzzling allele distributions as a result (Nichols and Hewitt 1994). Genetic drift is a process that will affect any pattern of genetic variation in a random fashion. Local adaptation through natural selection is a process that, if sufficiently strong in comparison with gene flow and genetic drift, will create patterns where genetic differentiation is associated with certain environmental conditions (Wright 1951). Several studies have shown the importance of local adaptation of populations (reviewed by Kawecki and Ebert 2004) (see also Bonin et al. 2006; Knight and Miller 2004; Kolseth and Lönn 2005; Lönn et al. 1998). Local adaptation can be strong also at small spatial scales (Snaydon and Davies 1976; Lönn 1993) even though it is sometimes very limited in terms of the number of genes involved (Kärkkainen et al. 2004) Environmental variability provides a base for biological variation by imposing differentiated selection pressures resulting in local adaptation. Topography provides large environmental variation within a relatively small area and thereby provides a basis for small-scale local adaptations. Depending on the local topographic possibilities populations can either migrate up and down slopes or along the same altitude to a different aspect to find a suitable microclimate. The dispersal distance will be much shorter per degree of temperature change during altitudinal migration (Hewitt 1996), than during simple latitudinal migration across a flat landscape. Slope and aspect are two important topographic parameters that determine the influx level of solar radiation, especially towards the poles where the total global radiation decreases (Larcher 2003). Vicia pisiformis is an endangered poorly-dispersed long-lived forest herb with its main distribution across the semi-open broadleaved forest steppes of eastern Europe. The Scandinavian populations are believed to be climate relict populations from warmer times. Earlier genetic studies of V. pisiformis using allozymes, RAPD:s and morhology, have found low to very low levels of genetic variation (Gustafsson and Gustafsson 1994; Black-Samuelsson et al. 1997; Black-Samuelsson and Lascoux 1999). Therefore we used AFLP (amplified fragment length polymorphism) markers, which detect even very small genetic differences between individuals. AFLP mainly analyse neutral variation, as the major fraction of most genomes is assumed to be neutral. However, since the AFLP-fragments are distributed randomly throughout the whole genome some fragments may be situated so close to regions under selection that they become more or less linked to them. This linkage disequilibrium between molecular markers and regions under selection, often referred to as quantitative trait loci (QTL), forms the basis for both QTL-mapping and marker assisted selection (MAS), reviewed by Dekkers and Hospital (2002). Gardner and Latta (2006) for example, found QTL under selection in both natural environments and in the greenhouse. Markers have been found to be connected to biomass production (Cavagnaro et al. 2006) and environmental variation (Bonin et al. 2006; Jump et al. 2006; Porcher et al. 2006). In this study we examine 22 Swedish and Norwegian populations of Vicia pisiformis and ask (i) if there is genetic differentiation between these populations, (ii) if there is can it be explained in its entirety by geographic location or (iii) can it partly be explained by habitat characteristics, suggesting local adaptation, or population size, suggesting genetic drift. We show that populations are differentiated geographically and that genetic variation in addition to the geographical pattern is associated with habitat.

Vicia pisiformis L. is a perennial leguminous plant with a main distribution in broadleaved forest-steppes of eastern Europe. The species is classified as endangered (EN) according to the IUCN red-lists in both Norway and Sweden, due to severe fragmentation, small population sizes and continuing population decline. The populations on the Scandinavian Peninsula constitute the northern limit of the species distribution and are mostly restricted to warm stony slopes with predominantly southern aspects. In this study we used the AFLP method, which is a high-resolution genetic fingerprint method. Samples were collected from 22 Scandinavian populations. The overall genetic structure was analysed in an AMOVA, in a Mantel test and through constrained correspondence analysis (CCA). The ordination scores representing non-geographic genetic divergence were extracted from the CCA and analysed in a linear model using habitat variables and population size as explanatory variables. We found (i) a strong geographic structure, (ii) significant genetic divergence between populations, (iii) that this genetic divergence remained significant even after removing the effect of geography in a partial CCA and (iv) that the remaining non-geographic part of genetic divergence (distance from the ordination centre) was associated with aspect, populations with a northern aspect were more genetically divergent. Aspect explains more variation than population size and is the only variable retained in the minimal adequate model. We suggest that local adaptation has caused this divergence from an expected geographical pattern of genetic variation. This explanation is further supported by the association between aspect and specific AFLP fragments. Many plant populations are relics of a different climate (Aguirre-Planter et al. 2000; Despres et al. 2002; Pico and Riba 2002). In response to long-term climate change, populations can either migrate towards a more favourable climate or adapt to the new conditions (delaVega 1996; Jump et al. 2006). Species with limited dispersal ability are at risk of reaching isolated dead-ends of decreasingly suitable habitat, without any suitable habitat within dispersal distance (Colas et al. 1997). Isolated populations have to use their inherent evolutionary potential and adapt to changes in environmental conditions, or they will go extinct. As population fragments go extinct, those that remain will become increasingly isolated from each other both spatially and also genetically as the level of gene flow declines with increasing distance. Such correlation between genetic dissimilarities and geographic distances, known as isolation by distance (Slatkin 1993; Wright 1943), when found, suggests a history of geographically limited gene flow (Kimura and Weiss 1964). On top of an isolation by distance pattern there might be other genetic structures to be found. Occasional long-distance dispersal events for example may disturb geographic patterns with puzzling allele distributions as a result (Nichols and Hewitt 1994). Genetic drift is a process that will affect any pattern of genetic variation in a random fashion. Local adaptation through natural selection is a process that, if sufficiently strong in comparison with gene flow and genetic drift, will create patterns where genetic differentiation is associated with certain environmental conditions (Wright 1951). Several studies have shown the importance of local adaptation of populations (reviewed by Kawecki and Ebert 2004) (see also Bonin et al. 2006; Knight and Miller 2004; Kolseth and Lönn 2005; Lönn et al. 1998). Local adaptation can be strong also at small spatial scales (Snaydon and Davies 1976; Lönn 1993) even though it is sometimes very limited in terms of the number of genes involved (Kärkkainen et al. 2004) Environmental variability provides a base for biological variation by imposing differentiated selection pressures resulting in local adaptation. Topography provides large environmental variation within a relatively small area and thereby provides a basis for small-scale local adaptations. Depending on the local topographic possibilities populations can either migrate up and down slopes or along the same altitude to a different aspect to find a suitable microclimate. The dispersal distance will be much shorter per degree of temperature change during altitudinal migration (Hewitt 1996), than during simple latitudinal migration across a flat landscape. Slope and aspect are two important topographic parameters that determine the influx level of solar radiation, especially towards the poles where the total global radiation decreases (Larcher 2003). Vicia pisiformis is an endangered poorly-dispersed long-lived forest herb with its main distribution across the semi-open broadleaved forest steppes of eastern Europe. The Scandinavian populations are believed to be climate relict populations from warmer times. Earlier genetic studies of V. pisiformis using allozymes, RAPD:s and morhology, have found low to very low levels of genetic variation (Gustafsson and Gustafsson 1994; Black-Samuelsson et al. 1997; Black-Samuelsson and Lascoux 1999). Therefore we used AFLP (amplified fragment length polymorphism) markers, which detect even very small genetic differences between individuals. AFLP mainly analyse neutral variation, as the major fraction of most genomes is assumed to be neutral. However, since the AFLP-fragments are distributed randomly throughout the whole genome some fragments may be situated so close to regions under selection that they become more or less linked to them. This linkage disequilibrium between molecular markers and regions under selection, often referred to as quantitative trait loci (QTL), forms the basis for both QTL-mapping and marker assisted selection (MAS), reviewed by Dekkers and Hospital (2002). Gardner and Latta (2006) for example, found QTL under selection in both natural environments and in the greenhouse. Markers have been found to be connected to biomass production (Cavagnaro et al. 2006) and environmental variation (Bonin et al. 2006; Jump et al. 2006; Porcher et al. 2006). In this study we examine 22 Swedish and Norwegian populations of Vicia pisiformis and ask (i) if there is genetic differentiation between these populations, (ii) if there is can it be explained in its entirety by geographic location or (iii) can it partly be explained by habitat characteristics, suggesting local adaptation, or population size, suggesting genetic drift. We show that populations are differentiated geographically and that genetic variation in addition to the geographical pattern is associated with habitat.

Genetic differentiation between and within five varieties of Euphrasia stricta (var. brevipila, var. gotlandica, var. stricta, var. suecica and var. tenuis) on Gotland was investigated, using amplified fragment length polymorphism, AFLP. The varieties are described in the literature by morphology and association to habitat type. We wanted to investigate whether the varieties are locally adapted populations to the typical habitat type for each variety or if they are preadapted to certain habitat types and have colonized Gotland in their present form. A constrained principal coordinate analysis revealed three genetically differentiated subunits within the species. The two early-flowering varieties suecica and tenuis each formed a distinct group, while the three late-flowering varieties brevipila, gotlandica and stricta together formed the third group. A phylogenetic tree confirms the partitioning into three groups. Within the group containing the late-flowering varieties there are populations that pair as each other's closest relatives, but belong to different varieties. These pairs are also geographically adjacent. The phylogenetic tree had a “star-like” appearance indicating a stronger divergence between populations than between varieties. The same pattern was seen in the partitioning of genetic diversity, with a lower amount of genetic variation occurring between varieties, F_ST=0.14, than between populations within the varieties, F_ST ranging from 0.26 to 0.60. In Euphrasia stricta the varieties suecica and tenuis and the group containing the varieties stricta/gotlandica/brevipila are likely to have a phylogeographical history outside Gotland, or an ancient and concealed local origin on the island. Within the group stricta/gotlandica/brevipila local evolutionary events seem to determine the variety identity, probably through local adaptation. Natural selection, genetic drift and mutations create genetic differentiation between populations. Gene flow, on the other hand, may counteract these processes (Slatkin 1987). Local adaptation is affected by the stability and strength of the natural selection and the amount of gene flow (Rice and Mack 1991, van Tienderen 1992, Miller and Fowler 1994), but also by the amount of genetic variability for the character that selection works on (Dudley 1996). Many studies have been done in the area of local adaptation (Lönn 1993, Prentice et al. 1995, Lönn et al. 1996, Liviero et al. 2002), and some studies have identified selective agents causing the adaptations. The selective agents are for example small differences in ecological niches and frequency dependent selection caused by pathogens (Parker 1994) or differences in selection regimes in different habitats (Kittelson and Maron 2001). Recent findings on ecological speciation emphasizes the importance of niche-shifting in local populations or groups of populations (Levin 2003) and rapid accumulation of beneficial mutations in isolated small populations (Rieseberg et al. 2003). Evolution works on different spatial and temporal scales, which makes it important to consider these different scales when studying evolutionary processes. Looking at the local phylogeny, geographic and temporal aspects are important when they link evolutionary processes to the extant landscape and the properties of the genetic structure. Linking evolutionary processes to the extant landscape is an important tool in evaluating evolutionary potential and predicting effects of landscape changes. Regional dynamics within species, using varieties/ecotypes, may also give information on initiation of speciation events. Manel et al. (2003) introduce and define landscape genetics as the combination of molecular population genetics and landscape ecology. The advantage of landscape genetics is the combination of the broad geographical span of scales (landscape to microclimate) and the high genetic resolution (individuals) compared to biogeography and phylogeography, which focuses more on species level at a broad spatial and temporal scale. Escudero et al. (2003), like Manel et al. (2003), put an emphasis on the spatial analysis of genetic diversity where a second step is to find ecological or demographic processes that could have shaped the genetic structure. A more direct approach is to measure habitat and genetic properties at many geographic locations and then model the biological processes shaping the spatial genetic structure (Lönn 1993, Prentice et al. 1995), which is the approach we intend to follow here. Molecular markers will be able to trace stochastic processes like drift and gene-flow (Page and Holmes 1998) as well as selective events through hitch-hiking (Hedrick 1980) and linkage events: AFLP has been used to identify quantitative trait loci by Via and Hawthorne (2002) and to explore the role of directional selection in whitefish ecotypes by Campbell and Bernatchez (2004). Yeo (1954, 1956, 1961, 1962, 1964, 1966, 1968) has done an extensive study of the cytology, hybridisation, cultivation, germination and relationship between species of British and European Euphrasia species. Yeo (1968) concludes that differences in chromosome number, habitat preferences and spatial distribution drives the speciation of Euphrasia in Europe and limits the hybridisation between species. The hybridisation may however result in new gene combinations for selection to work on (Yeo 1968). Yeo (1968) suggests that Euphrasia has gone through a fast and quite recent evolution in Europe after the last glaciation since Euphrasia has interfertile species of which many are endemic to small areas. Today, species differentiation within Euphrasia may be due to vegetation history, hybridisation and the parallel selection of well-adapted biotypes in similar or identical habitats (Karlsson 1976). Both Karlsson (1986) and Yeo (1968) put emphasis on the habitat specialization as an important factor in speciation referring to high morphological variability and hybridisation creating possibilities to evolve habitat specializations in Euphrasia.Zopfi (1998) showed in cultivation experiments that there is a genetic basis for different ecotypic variants of Euphrasiarostkoviana defined by grassland management, concerning onset of flowering, seed size and flowering period, life-history characters that are important adaptations to grazing and mowing. Euphrasia stricta is a tetraploid annual hemiparasite belonging to the Scrophulariaceae family (Yeo 1968, Krok and Almquist 2001). The species occurs all over Europe, except on the British Isles and in Spain and Portugal (Hultén and Fries 1986). In Sweden five varieties are found, which are subdivided based on morphology, phenology and habitat preference (Krok and Almquist 2001). They all grow on the Baltic island of Gotland, which is situated east of Sweden consisting of Silurian limestone (Fredén 1994). Euphrasia stricta var. suecica and E. stricta var. tenuis grow in traditionally managed wooded hay meadows and both are early flowering (Karlsson 1984). The variety suecica is red-listed according to the Swedish Red List (Gärdenfors 2000) and exists only in meadows on Gotland. The variety tenuis exists not only on Gotland but also on the Swedish mainland although it is declining throughout its distribution range. The populations of suecica and tenuis on Gotland are well known (Karlsson 1984, Petersson 1999). Euphrasia stricta var. stricta and E. stricta var. brevipila occur in pastures, along paths and on cultivated land. They flower later in the summer than var. suecica and var. tenuis. The variety stricta is common on the calcareous ground on Gotland with short grass turf, but rare on the mainland in contrast to the variety brevipila, which is common in whole of Sweden except on Gotland. The variety brevipila prefers soils that contain more sand compared to stricta. The late-flowering variety gotlandica is only found on Gotland and Öland, the second Baltic island on the Swedish east coast, were it is restricted to temporary pools on limestone ground (alvar) (Karlsson 1986). All Euphrasia species seems to lack a persistent seed bank (Karlsson 1984), but seeds have survived for up to three years in pots in cultivation experiments of other Euphrasia species (Yeo 1961). Artificial selfing and crosses within and between populations of Euphrasia stricta var. stricta yield high fertility in progeny pollen, 70–100% in between population crosses and 90–100% in selfing or within population crosses (Karlsson 1986). Flowering time for Euphrasia is not only dependent on habitat, but also to some extent on temperature and host attachment (Wilkins 1963, Yeo 1964, Molau 1993, Svensson et al. 2001, Svensson and Carlsson 2004). Euphrasia stricta probably have a mixed mating system (von Wettstein 1896). Based on these factors, which separates the varieties spatially and temporal, the aim of this study was to examine whether the varieties are locally adapted ecotypes that have evolved more than one time on the studied geographical scale or if they are distinct units over the region, implying colonization from outside or a single evolutionary event

Genetic differentiation between and within five varieties of Euphrasia stricta (var. brevipila, var. gotlandica, var. stricta, var. suecica and var. tenuis) on Gotland was investigated, using amplified fragment length polymorphism, AFLP. The varieties are described in the literature by morphology and association to habitat type. We wanted to investigate whether the varieties are locally adapted populations to the typical habitat type for each variety or if they are preadapted to certain habitat types and have colonized Gotland in their present form. A constrained principal coordinate analysis revealed three genetically differentiated subunits within the species. The two early-flowering varieties suecica and tenuis each formed a distinct group, while the three late-flowering varieties brevipila, gotlandica and stricta together formed the third group. A phylogenetic tree confirms the partitioning into three groups. Within the group containing the late-flowering varieties there are populations that pair as each other's closest relatives, but belong to different varieties. These pairs are also geographically adjacent. The phylogenetic tree had a “star-like” appearance indicating a stronger divergence between populations than between varieties. The same pattern was seen in the partitioning of genetic diversity, with a lower amount of genetic variation occurring between varieties, F_ST=0.14, than between populations within the varieties, F_ST ranging from 0.26 to 0.60. In Euphrasia stricta the varieties suecica and tenuis and the group containing the varieties stricta/gotlandica/brevipila are likely to have a phylogeographical history outside Gotland, or an ancient and concealed local origin on the island. Within the group stricta/gotlandica/brevipila local evolutionary events seem to determine the variety identity, probably through local adaptation. Natural selection, genetic drift and mutations create genetic differentiation between populations. Gene flow, on the other hand, may counteract these processes (Slatkin 1987). Local adaptation is affected by the stability and strength of the natural selection and the amount of gene flow (Rice and Mack 1991, van Tienderen 1992, Miller and Fowler 1994), but also by the amount of genetic variability for the character that selection works on (Dudley 1996). Many studies have been done in the area of local adaptation (Lönn 1993, Prentice et al. 1995, Lönn et al. 1996, Liviero et al. 2002), and some studies have identified selective agents causing the adaptations. The selective agents are for example small differences in ecological niches and frequency dependent selection caused by pathogens (Parker 1994) or differences in selection regimes in different habitats (Kittelson and Maron 2001). Recent findings on ecological speciation emphasizes the importance of niche-shifting in local populations or groups of populations (Levin 2003) and rapid accumulation of beneficial mutations in isolated small populations (Rieseberg et al. 2003). Evolution works on different spatial and temporal scales, which makes it important to consider these different scales when studying evolutionary processes. Looking at the local phylogeny, geographic and temporal aspects are important when they link evolutionary processes to the extant landscape and the properties of the genetic structure. Linking evolutionary processes to the extant landscape is an important tool in evaluating evolutionary potential and predicting effects of landscape changes. Regional dynamics within species, using varieties/ecotypes, may also give information on initiation of speciation events. Manel et al. (2003) introduce and define landscape genetics as the combination of molecular population genetics and landscape ecology. The advantage of landscape genetics is the combination of the broad geographical span of scales (landscape to microclimate) and the high genetic resolution (individuals) compared to biogeography and phylogeography, which focuses more on species level at a broad spatial and temporal scale. Escudero et al. (2003), like Manel et al. (2003), put an emphasis on the spatial analysis of genetic diversity where a second step is to find ecological or demographic processes that could have shaped the genetic structure. A more direct approach is to measure habitat and genetic properties at many geographic locations and then model the biological processes shaping the spatial genetic structure (Lönn 1993, Prentice et al. 1995), which is the approach we intend to follow here. Molecular markers will be able to trace stochastic processes like drift and gene-flow (Page and Holmes 1998) as well as selective events through hitch-hiking (Hedrick 1980) and linkage events: AFLP has been used to identify quantitative trait loci by Via and Hawthorne (2002) and to explore the role of directional selection in whitefish ecotypes by Campbell and Bernatchez (2004). Yeo (1954, 1956, 1961, 1962, 1964, 1966, 1968) has done an extensive study of the cytology, hybridisation, cultivation, germination and relationship between species of British and European Euphrasia species. Yeo (1968) concludes that differences in chromosome number, habitat preferences and spatial distribution drives the speciation of Euphrasia in Europe and limits the hybridisation between species. The hybridisation may however result in new gene combinations for selection to work on (Yeo 1968). Yeo (1968) suggests that Euphrasia has gone through a fast and quite recent evolution in Europe after the last glaciation since Euphrasia has interfertile species of which many are endemic to small areas. Today, species differentiation within Euphrasia may be due to vegetation history, hybridisation and the parallel selection of well-adapted biotypes in similar or identical habitats (Karlsson 1976). Both Karlsson (1986) and Yeo (1968) put emphasis on the habitat specialization as an important factor in speciation referring to high morphological variability and hybridisation creating possibilities to evolve habitat specializations in Euphrasia.Zopfi (1998) showed in cultivation experiments that there is a genetic basis for different ecotypic variants of Euphrasiarostkoviana defined by grassland management, concerning onset of flowering, seed size and flowering period, life-history characters that are important adaptations to grazing and mowing. Euphrasia stricta is a tetraploid annual hemiparasite belonging to the Scrophulariaceae family (Yeo 1968, Krok and Almquist 2001). The species occurs all over Europe, except on the British Isles and in Spain and Portugal (Hultén and Fries 1986). In Sweden five varieties are found, which are subdivided based on morphology, phenology and habitat preference (Krok and Almquist 2001). They all grow on the Baltic island of Gotland, which is situated east of Sweden consisting of Silurian limestone (Fredén 1994). Euphrasia stricta var. suecica and E. stricta var. tenuis grow in traditionally managed wooded hay meadows and both are early flowering (Karlsson 1984). The variety suecica is red-listed according to the Swedish Red List (Gärdenfors 2000) and exists only in meadows on Gotland. The variety tenuis exists not only on Gotland but also on the Swedish mainland although it is declining throughout its distribution range. The populations of suecica and tenuis on Gotland are well known (Karlsson 1984, Petersson 1999). Euphrasia stricta var. stricta and E. stricta var. brevipila occur in pastures, along paths and on cultivated land. They flower later in the summer than var. suecica and var. tenuis. The variety stricta is common on the calcareous ground on Gotland with short grass turf, but rare on the mainland in contrast to the variety brevipila, which is common in whole of Sweden except on Gotland. The variety brevipila prefers soils that contain more sand compared to stricta. The late-flowering variety gotlandica is only found on Gotland and Öland, the second Baltic island on the Swedish east coast, were it is restricted to temporary pools on limestone ground (alvar) (Karlsson 1986). All Euphrasia species seems to lack a persistent seed bank (Karlsson 1984), but seeds have survived for up to three years in pots in cultivation experiments of other Euphrasia species (Yeo 1961). Artificial selfing and crosses within and between populations of Euphrasia stricta var. stricta yield high fertility in progeny pollen, 70–100% in between population crosses and 90–100% in selfing or within population crosses (Karlsson 1986). Flowering time for Euphrasia is not only dependent on habitat, but also to some extent on temperature and host attachment (Wilkins 1963, Yeo 1964, Molau 1993, Svensson et al. 2001, Svensson and Carlsson 2004). Euphrasia stricta probably have a mixed mating system (von Wettstein 1896). Based on these factors, which separates the varieties spatially and temporal, the aim of this study was to examine whether the varieties are locally adapted ecotypes that have evolved more than one time on the studied geographical scale or if they are distinct units over the region, implying colonization from outside or a single evolutionary event

Connectivity plays an important role in shaping the genetic structure and in evolution of local adaptation. In the marine environment barriers to gene flow are in most cases caused by gradients in environmental factors, ocean circulation and/or larval behavior. Despite the long pelagic larval stages, with high potential for dispersal many marine organisms have been shown to have a fine scale genetic structuring. In this study, by using a combination of high-resolution genetic markers, species hybridization data and biophysical modeling we can present a comprehensive picture of the evolutionary landscape for a keystone species in the Baltic Sea, the blue mussel. We identified distinct genetic differentiation between the West Coast, Baltic Proper and Bothnian Sea regions, with lower gene diversity in the Bothnian Sea. Oceanographic connectivity together with salinity and to some extent species identity provides explanations for the genetic differentiation between the West Coast and the Baltic Sea (Baltic Proper and Bothnian Sea). The genetic differentiation between the Baltic Proper and Bothnian Sea cannot be directly explained by oceanographic connectivity, species identity or salinity, while the lower connectivity to the Bothnian Sea may explain the lower gene diversity. © 2016.

Connectivity plays an important role in shaping the genetic structure and in evolution of local adaptation. In the marine environment barriers to gene flow are in most cases caused by gradients in environmental factors, ocean circulation and/or larval behavior. Despite the long pelagic larval stages, with high potential for dispersal many marine organisms have been shown to have a fine scale genetic structuring. In this study, by using a combination of high-resolution genetic markers, species hybridization data and biophysical modeling we can present a comprehensive picture of the evolutionary landscape for a keystone species in the Baltic Sea, the blue mussel. We identified distinct genetic differentiation between the West Coast, Baltic Proper and Bothnian Sea regions, with lower gene diversity in the Bothnian Sea. Oceanographic connectivity together with salinity and to some extent species identity provides explanations for the genetic differentiation between the West Coast and the Baltic Sea (Baltic Proper and Bothnian Sea). The genetic differentiation between the Baltic Proper and Bothnian Sea cannot be directly explained by oceanographic connectivity, species identity or salinity, while the lower connectivity to the Bothnian Sea may explain the lower gene diversity. © 2016.

Human-derived environmental pollutants and nutrients that reach the aquatic environment through sewage effluents, agricultural and industrial processes are constantly contributing to environmental changes that serve as drivers for adaptive responses and evolutionary changes in many taxa. In this study, we examined how two types of point sources of aquatic environmental pollution, harbors and sewage treatment plants, affect gene diversity and genetic differentiation in the blue mussel in the Baltic Sea area and off the Swedish west coast (Skagerrak). Reference sites (REF) were geographically paired with sites from sewage treatments plant (STP) and harbors (HAR) with a nested sampling scheme, and genetic differentiation was evaluated using a high-resolution marker amplified fragment length polymorphism (AFLP). This study showed that genetic composition in the Baltic Sea blue mussel was associated with exposure to sewage treatment plant effluents. In addition, mussel populations from harbors were genetically divergent, in contrast to the sewage treatment plant populations, suggesting that there is an effect of pollution from harbors but that the direction is divergent and site specific, while the pollution effect from sewage treatment plants on the genetic composition of blue mussel populations acts in the same direction in the investigated sites.

Human-derived environmental pollutants and nutrients that reach the aquatic environment through sewage effluents, agricultural and industrial processes are constantly contributing to environmental changes that serve as drivers for adaptive responses and evolutionary changes in many taxa. In this study, we examined how two types of point sources of aquatic environmental pollution, harbors and sewage treatment plants, affect gene diversity and genetic differentiation in the blue mussel in the Baltic Sea area and off the Swedish west coast (Skagerrak). Reference sites (REF) were geographically paired with sites from sewage treatments plant (STP) and harbors (HAR) with a nested sampling scheme, and genetic differentiation was evaluated using a high-resolution marker amplified fragment length polymorphism (AFLP). This study showed that genetic composition in the Baltic Sea blue mussel was associated with exposure to sewage treatment plant effluents. In addition, mussel populations from harbors were genetically divergent, in contrast to the sewage treatment plant populations, suggesting that there is an effect of pollution from harbors but that the direction is divergent and site specific, while the pollution effect from sewage treatment plants on the genetic composition of blue mussel populations acts in the same direction in the investigated sites.

We have recently found that the morphologically determined subspecies Gymnadenia conopsea ssp conopsea in Sweden includes early and late flowering individuals. We were interested in the interactions between the flowering time groups; if there were gene flow between them and if so this was detrimental or advantageous. A spatially mixed population of early and late flowering individuals was studied using three microsatellite loci. We measured patterns in genetic differentiation and inferred occurrence of hybridisation and introgression. Variation in flowering time, fertility and relative and absolute fruit set was measured. The pattern of introgression between flowering-time groups differed between loci. In two of the three investigated loci, allele separation was distinct between early and late flowering plants and one genetically obvious hybrid was infertile. In the third locus, several alleles were shared between the two flowering time variants. The degree of introgression was associated to fruit set failure, which was higher in the late flowering plants and lower in early flowering plants. A small group of early flowering individuals with somewhat delayed flowering compared to the main group was genetically distinct and had lower relative and absolute fruit set. This group was not genetically intermediate, but rather constituting an independent group, with lower fruit set possibly caused by absence of pollinators. There seem to be a strong barrier against introgression into the late flowering group which is kept genetically distinct and less diverse. The early flowering group is diverse, includes two subgroups and seems to benefit from gene flow

We have recently found that the morphologically determined subspecies Gymnadenia conopsea ssp conopsea in Sweden includes early and late flowering individuals. We were interested in the interactions between the flowering time groups; if there were gene flow between them and if so this was detrimental or advantageous. A spatially mixed population of early and late flowering individuals was studied using three microsatellite loci. We measured patterns in genetic differentiation and inferred occurrence of hybridisation and introgression. Variation in flowering time, fertility and relative and absolute fruit set was measured. The pattern of introgression between flowering-time groups differed between loci. In two of the three investigated loci, allele separation was distinct between early and late flowering plants and one genetically obvious hybrid was infertile. In the third locus, several alleles were shared between the two flowering time variants. The degree of introgression was associated to fruit set failure, which was higher in the late flowering plants and lower in early flowering plants. A small group of early flowering individuals with somewhat delayed flowering compared to the main group was genetically distinct and had lower relative and absolute fruit set. This group was not genetically intermediate, but rather constituting an independent group, with lower fruit set possibly caused by absence of pollinators. There seem to be a strong barrier against introgression into the late flowering group which is kept genetically distinct and less diverse. The early flowering group is diverse, includes two subgroups and seems to benefit from gene flow