This thesis is a summary of enclosure and microcosm experiments that aimed to study the impact of allochtonous subsidies on food web dynamics in a heterogeneous food web. In the enclosure studies, a three trophic level littoral food web was used. The food web consisted of two growth forms of primary producers: phytoplankton and periphyton and their associated herbivores: scraping and filtering herbivores. The predator used, YOY perch, affects both pathways in the food web. Manipulation of the openness for the different trophic levels showed that the inflow of phytoplankton and cross-habitat foraging by the herbivore level reinforced the compensatory response between the two growth forms of primary producers In the microcosm experiment, the response of an herbivore food web and a microbial community to inflow of resources and food web configuration was studied, using a model food web. The model food web consisted of two forms of primary producers, i.e. phytoplankton and periphyton, and two herbivores, i.e. Daphnia pulex feeding on phytoplankton, and Chydorus sphaericus feeding on both periphyton and phytoplankton. Three different food web configurations, all having the phytoplankton and periphyton, but either one of the herbivores, or both, were set up. The flow regimes consisted of an open treatment receiving a constant supply of phytoplankton, and a closed treatment with an initial resource pool. The effect of the inflow of phytoplankton was affected by the food web configuration. In the presence of D. pulex, the inflow of phytoplankton was made accessible to periphyton, and indirectly to C. sphaericus, which increased to such high densities that D. pulex was negatively affected. The inflow of phytoplankton had an indirect negative effect on the microbial community, since the biomass of herbivores increased, which imposed a higher grazing pressure on all parts of the microbial community.

The main objectives of this thesis are to study if a topographic wetness index (TWI) could be used as a tool for predicting the spatial distribution of vascular plant species richness in the boreal forest as well as to study congruence in species richness between vascular plants, liverworts, mosses and lichens. A wetness index ln(a/tanβ) based on topography was used to assign a specific TWI-value to every 20 x 20m grid in two 25 km2 boreal forest landscapes (differing in average soil pH) in northern Sweden. Soil pH is known to be influenced by groundwater and to affect plant species richness in other biomes. Therefore, the relationships between plant species richness, TWI and soil pH were also studied. The results showed that the majority of the investigated boreal forest landscapes were relatively dry and species-poor, whereas interspersed patches linked to areas with relatively high TWI had species-rich vegetation including the species of the drier parts of the landscape. There was a positive relationship between species richness of vascular plants and the TWI in both landscapes, but varied with average soil pH. TWI explained 30 % and 52 % of the variation in plant species richness in the landscape with lower and higher pH, respectively. The proportion of regionally uncommon plants also increased with TWI. Testing different calculation methods of the TWI resulted in a large variation in correlation strengths between the various TWI-values and different measured variables (species richness of vascular plants, soil pH, groundwater flow and soil moisture). The relationship between plant species richness and TWI could be further improved with some of the calculation methods. When studying correlations in species richness using data sets from boreal forest in northern Sweden, strong positive correlations among vascular plants, mosses and liverworts were found, but no significant correlation between macrolichens and any of the other groups. This result could be explained by that the species number of each of the three related groups increases with ambient moisture, whereas the species number of macrolichens is weakly associated with moisture. In conclusion, the TWI could become a useful tool in conservation management for identifying areas of special interest prior to field inventories. Since vascular plants can be used as an indicator taxon for species richness of mosses and liverworts, high TWI-values indicate areas of species hotspots of these taxa. / Syftet med avhandlingen är att dels studera om ett topografiskt fuktighetsindex skulle kunna vara användbart för att förutsäga fördelningen av kärlväxters artrikedom i boreal skog, dels att studera om den rumsliga fördelningen av artrikedom hos kärlväxter, blad- och levermossor samt lavar sammanfaller. Ett fuktighetsindex, ln(a/tanβ), som bara är baserat på topografi användes för att beräkna ett indexvärde för varje 20 x 20 m grid i två 25 km2 stora boreala skogslandskap (med i medeltal olika mark-pH) i norra Sverige. Det är känt att mark-pH påverkas av grundvatten och att pH i sin tur påverkar artrikedom hos kärlväxter i andra biom. Därför studerades även sambanden mellan kärlväxters artrikedom, mark pH och TWI. Resultaten visade att större delen av det studerade boreala landskapet var relativt torrt och artfattigt, medan mindre utspridda områden med höga TWI-värden var artrika på kärlväxter och här växte även arter som återfanns i de torra delarna av skogen. Sambandet mellan artrikedom hos kärlväxter och TWI var positivt i båda landskapen, men påverkades av de olika nivåerna på mark-pH. TWI förklarade 30 % av variationen i artrikedom i området med lägre mark-pH respektive 50 % i området med högre mark-pH. Andelen kärlväxter som klassificeras som icke vanliga i respektive region ökade också med TWI. Med andra beräkningsmetoder för TWI visade det sig att styrkan på korrelationerna mellan TWI och olika uppmätta variabler (artrikedom hos kärlväxter, mark-pH, grundvattennivå och markfuktighet) varierade mycket. Sambandet mellan artrikedom hos kärlväxter och TWI kunde förbättras ytterligare med vissa beräkningsmetoder. Då korrelationer i artrikedom studerades användes ett dataset från boreal skog i norra Sverige. Resultaten visade på starka, positiva korrelationer mellan kärlväxter, blad- och levermossor, men inga korrelationer mellan någon av dessa grupper och lavar. Detta kunde förklaras med att artrikedom hos de tre korrelerande organismgrupperna ökar med ökad fuktighet, medan artrikedom hos lavar inte är kopplat till fukt. Huvudslutsatsen i avhandlingen är att TWI, som endast är baserad på topograpfiskt data, skulle kunna bli ett värdefullt redskap i naturvårdsplanering för att identifiera särskilt intressanta skogsområden innan man gör fältinventeringar. Eftersom studien visar att kärlväxter kan användas som en indikator grupp för artrikedom hos blad- och levermossor indikerar höga TWI-värden områden med hög artrikedom även vad gäller dessa taxa.

Sexual selection arises through variation in reproductive success. This thesis investigates different aspects important in sexual selection, namely nest building, sperm competition, paternity and paternal care, and their mutual interrelationships. In the studied species, the sand goby (Pomatoschistus minutus) and the common goby (Pomatoschistus microps), sperm competition did arise when small males, so called sneakers, sneaked into other males nests and released sperm. They seemed to use female behaviour as their prime cue for a sneaking opportunity. However, also nest-holders, both with and without eggs, were found to fertilize eggs in the nests of other males. Clearly, nest-holding males tried to prevent other males from spreading their sperm in their nests, since they showed aggression towards such males. A nest building experiment indicated that the small nest-openings found in the sneaker male treatment were sexually selected through protection against sneaking or by female choice. Yet, no behavioural or genetical support for the hypothesis that the nest functions as a physical or visual defence, or that sneaker males prefer to sneak upon nests with wide nest-openings, were found in the other studies. Still, individual nest-holding males showed a higher mucus preparation effort inside the nest in the presence of a sneaker male than when alone. In close relatives, such mucus contains sperm, suggesting an importance in sperm competition. However, the mucus may also have pheromone and anti-bacterial functions and may constitute a mating effort, as found in other gobies. Both a behavioural and a mate choice experiment suggested that the males were not less eager to spawn in the presence of a sneaker male. Sneak intrusion did not affect nest defence, fanning or filial cannibalism, nor had paternity an effect on filial cannibalism. This and various life history aspects, together with the fact that the parasitic male only fertilized a fraction of the clutches, would predict females to ignore sneaker males. This was also the case, as the presence of sneaker males was found not to affect female spawning decision. Still, several females spawned in two nests, which coincided with parasitic spawnings, suggesting a cost of disturbance for the females and thus a substantial cost to the nest-holding males in terms of lost mating success. However, females paid attention to other traits in their choice of mate since spawning was associated with sand volume of the nest, but not with nest-opening width. Also, female (but not male) courtship was correlated with partial clutch filial cannibalism, indicating that females are able to anticipate future male cannibalism. In a partial correlation of nest opening, sand volume, male courtship display, displacement fanning and male size, a large number of traits were correlated both positively and negatively with regard to how we may expect them to be appreciated by females. For instance, males which fan well also build large nests or display intensely (but not both). Together with all the other results of this thesis, this shows the entangled selection pressures working on breeding animals, as well as the different male and female tactics employed to maximize their reproduction.

The abundance of predators and their impact on ecosystem dynamics is a vividly discussed topic in current ecology. In my studies, incorporating field observations, field experiments and theoretical modeling, I explored the importance of predators and predation in a low arctic tundra ecosystem in northern Norway. This involved observing the abundance and spatial activity of predators (raptors and small mustelids); manipulating the abundance of predators (spiders and birds) in an arthropod community; and exploring the theoretical consequences of intraguild predation on the coexistence among predators. The results show that predation is important both in the arthropod assemblage and, depending on the productivity of the community, in the vertebrate assemblage. In arthropod communities predators are at least as abundant as their prey, whereas in the vertebrate part of ecosystem, predators are substantially less abundant than their prey. Still, in both cases predators had strong impact on their prey, influencing the abundance of prey and the species composition of prey assemblages. The impact of predation cascaded to the plant community both in the reticulate and complex arthropod food web and in the linear food chain-like vertebrate community. In the vertebrate-based community we could even observe the long time scale effect on plant community composition. Within the predator community, exploitation competition and intraguild predation were the structuring forces. As the arthropod communities consist of predators with different sizes, intraguild predation is an energetically important interaction for top predators. As a consequence, they reduce the abundance of intermediate predators and the impact of intermediate predators on other prey groups. Moreover, being supported by intermediate predators, top predators can have stronger impact themselves on other prey groups. In vertebrate communities, intraguild predation seems to be unimportant as energetic link, instead it manifests as an extreme version of interference competition. Therefore intraguild predation reduces the likelihood of coexistence, as it is due limited prey diversity and intense exploitative competition already precarious in the low arctic tundra. In conclusion, predators have strong impact on their prey, especially in the more productive parts of the low arctic tundra. This applies even to the food webs with complex and reticulate structure, and these effects carry through the community both in the short time scale of population growth and on the long time scale of population generations.

In this thesis, quantitative genetic approaches, common-garden experiments, and field studies were combined to examine patterns of population differentiation and the genetic architecture of characters of putative adaptive significance in the widely distributed perennial herb Lythrum salicaria. In this work, I (1) documented patterns of population differentiation in phenology, life-history, and morphology along latitudinal gradients at different geographical scales, (2) investigated the genetic architecture of phenology, flower morphology, and inflorescence size, and (3) combined estimates of phenotypic selection in the field with information on the genetic variance-covariance matrix (G) to examine potential constraints to adaptive evolution. A common-garden experiment demonstrated latitudinal variation in life-history, and phenology of growth and reproduction among L. salicaria populations sampled across Sweden. Flower morphology varied significantly among populations, but was, with the exception of calyx length, not related to latitude of origin. A second experiment, which included two Swedish, two Dutch, and two Spanish populations, indicated that the latitudinal gradient in reproductive and vegetative phenology might extend throughout Europe. A quantitative-genetic study of two Swedish populations revealed significant additive genetic variation for all phenological and morphological traits investigated. The G matrices of the populations differed significantly according to common principal component analysis, and genetic correlations within the study populations did not strictly correspond to trait correlations observed among populations. In a field study, I detected directional selection through female function for larger inflorescences in two consecutive years. Relative fitness increased disproportionately with inflorescence size in the year when supplemental hand-pollination indicated that pollen limitation was severe. Genetic correlations with inflorescence size considerably influenced predicted response to selection in other characters. Taken together, the results suggest that among-population differences in phenology and life-history in L. salicaria have evolved in response to latitudinal variation in length of the growing season. They demonstrate that the evolutionary potential of local populations may be considerable. The genetic covariance structure substantially influences predicted short-term evolutionary trajectories. However, the weak correspondence between genetic correlations documented within populations and trait correlations among populations, suggest that the G matrix has not imposed strict constraints on patterns of among-population differentiation.

The brackish Baltic Sea is a marginal environment for both marine and freshwater species. The rate of ecological differentiation is presumably high due to strong selection pressure from a gradient of decreasing salinity that has been present in its current state for only about 3 000 years. Even more recently, increased nutrient loading due to human activities has affected the growth rate of species, with potential effects on their competitive interactions and responses to other regulating factors. I have investigated the potential effects of low salinity and nutrient enrichment on the distributional ranges of two marine macroalgae with a wide distribution in the Baltic Sea, the red alga Ceramium tenuicorne (Kütz.) Wærn and the brown alga Fucus vesiculosus L. A field study in the northern Baltic Sea indicated a strong relationship between the community structure of macroalgae and abiotic factors even on a small, local scale. The abiotic factors are potentially modulated by eutrophication, which may have a strong effect on the depth distribution and abundance of macroalgae. On a regional scale, laboratory experiments suggested that nutrient enrichment is unlikely to affect the distribution of Ceramium and Fucus along the salinity gradient. Growth in Ceramium from the Baltic Proper was enhanced by nitrate and phosphate, but the response did not override growth constraints due to low salinity. Ceramium from the Gulf of Bothnia had an inherently lower growth rate that was not positively affected by nitrate and phosphate increase. In Fucus vesiculosus, reproductive performance was impaired by nitrate and phosphate levels corresponding to ambient levels in eutrophicated areas of the Baltic Sea, when measured by their effect on zygote attachment, germination, and rhizoid development. The wide distribution of Ceramium in the inner Baltic Sea is probably related to local adaptation, rather than a generalized tolerance of different salinity levels. Ecotypic differences were observed when comparing strains from the Baltic Proper (salinity 7 psu) and the Gulf of Bothnia (4 psu). A high rate of vegetative reproduction was evident, although sexual reproduction was occasionally observed in salinity 4. In Fucus vesiculosus, genetic and morphological analyses of sympatric and allopatric populations of the common, vesicular, morphotype and a dwarf morphotype, characteristic for the Gulf of Bothnia, showed that the dwarf morphotype represents a separate evolutionary lineage. Also, vegetative reproduction was observed in Fucus for the first time, as supported by genetic and experimental data. The results show that the biota of the inner Baltic Sea may have unique adaptive and genetic properties, and that it is highly relevant to consider subspecies diversity in Baltic Sea management.

Heterotrophic bacteria utilize dissolved organic matter, and the carbon flow through an ecosystem depends on the fractions of the utilized carbon that is either respired or transferred to higher trophic levels. The major aim this thesis is to investigate 1) the relationship between composition and functioning in heterotrophic bacterioplankton communities and 2) the influence of environmental conditions on both parameters. I set up several batch culture experiments, where lake water filtrates containing bacteria but no grazers were inoculated into sterile freshwater medium to investigate the importance of the origin of the source community (the inoculum) versus the environmental conditions (the medium) for the composition and functional performance of bacterial communities. In some experiments the medium was manipulated to simulate changes in salinity, pH and dissolved organic matter quantity and quality. Functional parameters (biomass yield, respiration, growth efficiency and enzyme activities) and the genetic composition of the emerging bacterial communities were determined. When bacterial inocula obtained from different habitats were re-grown under identical conditions, differently composed communities emerged. This indicates that the history and distribution of taxa within the inoculum was an important regulating factor of community composition. The coupling between community composition and functioning was not very tight, and there was functional equivalency with respect to aggregated functions important at the ecosystem scale (e.g., biomass production and respiration). The functional performance of bacterial communities could to a large extent be predicted from the medium alone, except when it deviated strongly from the ambient settings. When bacterial communities were exposed to dilution, a strong change in pH or an increase in salinity, growth of structurally and functionally distinct communities occurred. I therefore suggest that it depends on the disturbance regime how bacterial community structure and function are related to each other.

Bacteria and fungi dominate the decomposition of aquatic plants, a major process in the carbon and nutrient cycling in many aquatic systems. Although phylogenetically distant, bacteria and fungi often live in close proximity with each other. Since these microorganisms also have similar ecological functions, interactions have developed between them. This thesis explores the nature of such interactions, and the potential effects on key components of the decomposition process. The thesis includes a critical assessment of the ergosterol method for determination of fungal biomass, a survey of the environmental factors determining the distribution and taxa numbers of litter-decomposing bacteria and fungi in lakes, and a number of experiments on the interactions between bacteria and fungi. In all the experiments performed, fungi responded to bacterial presence through antagonism, although different fungal strains, bacterial communities and substrates were used. The antagonism seemed to be caused by interference competition for substrate. The fungal effect on bacteria was less consistent. Bacterial growth was suppressed, unaffected, or even enhanced by the presence of fungi. Fungi contributed more to extracellular enzyme production than bacteria, and bacteria were probably able to assimilate intermediate decomposition products formed through the activity of extracellular enzymes of fungal origin. Thus, the effect on bacteria from interacting with fungi was determined by the balance between competition and benefit from excreted enzymes. Bacteria and fungi also used different size fractions of the organic matter, according to their different enzymatic capacities. Hence, bacteria appeared to assimilate low-molecular-weight compounds, while high-molecular-weight compounds were utilized primarily by fungi. In brief, the ecological interactions influenced the growth and hence also the biomass development of bacteria and fungi, which affected enzyme activity as well as utilization of dissolved organic matter. Therefore, I suggest that interactions between bacteria and fungi influence degradation of plant litter in aquatic systems.

I combined comparative and experimental studies in the field and in the greenhouse to examine factors influencing reproductive success in two insect-pollinated herbs, the tristylous, selfincompatible perennial Lythrum salicaria and the self-compatible, biennial Pedicularis palustris. More specifically, I explored (i) the effects of plant population characteristics on the intensity and outcome of interactions with pollinators and seed predators, (ii) whether flower morphology affects duration of stigma receptivity, and (iii) whether damage-induced reduction in floral display reduces pollinator visitation and increases pollen limitation. As predicted, the rate of pollinator visitation tended to increase and the number of flowers probed per plant tended to decrease with increasing population size in L. salicaria, but these relationships only approached statistical significance. By taking advantage of the pollen size polymorphism that is typical of many heterostylous plants, I could show that the number of compatible pollen grains received increased with population size, and that this was associated with a reduction in pollen limitation and increased seed output per flower. The deposition of high numbers of incompatible conspecific and heterospecific pollen grains did not appear to reduce seed set. In P. palustris, fruit set and seed predation varied markedly among populations and years, but this variation could only partly be explained by variation in population size. Fruit set was positively related to population size, and seed predation was negatively related to population size, in one of three years. Similarly, the level of pollen limitation, which was quantified in two years, varied among populations, but was not related to population size, density or isolation. In L. salicaria, both the duration of stigma receptivity and the effect of prior self-pollination on seed output varied among style morphs. These differences may contribute to morph-specific differences in pollen limitation and seed production documented in the field. The results of a field experiment demonstrated that damage to the shoot apex may markedly reduce fruit production in L. salicaria, and suggested that this is mainly because damage reduces flower number. I found no evidence that a reduced floral display increased the severity of pollen limitation. Taken together, the results show that interactions with both pollinators and herbivores may markedly affect reproductive output in the two plant species, and that the intensity of both mutualistic and antagonistic interactions vary considerably in space and time. Moreover, they are consistent with the hypothesis that pollination success should depend less on population size in selfcompatible than in self-incompatible plants.

An influence at one trophic level can result in dynamic impacts also on other components of a food web. These dynamics are known as trophic cascades, and can be both top-down and bottom-up. After a near-collapse of the Baltic cod Gadus morhua stock in the 1980s, its main prey sprat Sprattus sprattus increased dramatically. The main food of sprat, marine copepods, decreased during the same time period, likely a combined effect of increased predation pressure from sprat and decreasing salinities. This shortage of food for sprat resulted in decreasing quality of sprat as a food source for common guillemots Uria aalge. However, a recent increase in fishing for sprat has again resulted in better feeding conditions for guillemots. Human impacts on this simple food web can be complex. In the early 20th century, marine mammals were abundant and nutrient levels were low in the Baltic Sea. This thesis illustrate that this situation corresponded to lower fish biomass. A reduction of seals early in the century led to reduced top-down control, which resulted in increasing fish stocks. Later, in the 1950s, the largest inflow of salt water during the century mobilized accumulated phosphorus from the deep sediments, which stimulated nitrogen fixation. Combined with increasing anthropogenic nutrient loads, this led to increased primary production and a rapid change from an oligotrophic to a eutrophicated state. This change can be termed a regime shift, which also stimulated fish production. Subsequent over-fishing of cod likely caused a second regime shift, from a cod- to a clupeid- dominated state, which led to the described effects on the common guillemots. Several factors affect the life-history of this long-lived seabird. Bycatches in gillnet fisheries is one factor directly affecting guillemot survival, and the proportion of bycatches increased during a period of increasing fishing effort. Surprisingly, avian cholera, a previously undocumented disease in common guillemots, was found at times to cause considerable adult mortality. Common guillemot life-history information can communicate the diversity of factors influencing marine ecosystems – hopefully this can increase our understanding of how complex even "simple" food webs are.