In this thesis I examine and evaluate the effects of a documented large-scale eutrophication on macroalgal vegetation on the Swedish coast. During the past century the load of nutrients has increased manifold in the Baltic Sea area, increasing primary production and organic sedimentation significantly. By re-investigating a unique reference material of macroalgal vegetation from the 1940-60s and by conducting new diving studies at the same sites, I showed that long-term trends in macroalgal community composition and species’ distributions are consistent with changes expected from an increased nutrient availability. In the Öregrund archipelago (northern Baltic Sea), I documented a declined depth distribution of the dominant canopy forming alga, Fucus vesiculosus, and an increased abundance of opportunistic ephemeral algae compared to 52-53 years ago. In the Gullmar Fjord area (Skagerrak), trends included increased abundances of functional groups with opportunistic algae, decreased abundances of large perennial algae and a general decline in the depth distribution of the vegetation compared to 36-57 years ago. Sediment removal experiments in the northern Baltic Sea confirmed the hypothesis that sedimentation influences macroalgal community composition. Species depending on short periods of reproduction were clearly favoured by sediment removal, especially F. vesiculosus that seemed limited in depth by the local sediment load. Species with long continuous periods of dispersal by spores and/or fragments (for example the ephemerals Cladophora glomerata, Ceramium tenuicorne and Enteromorpha spp.) were more tolerant to the natural sediment load. In general, sediment removal favoured macroalgal establishment and development, indicating that variation in the natural sediment load is an important constraint for sublittoral rocky-shore macroalgal community development. I conclude by suggesting that the documented long-term changes in macroalgal vegetation on the Swedish coast partly are explained by an increased organic sedimentation in these areas.

The main aim of this work was to identify local adaptation processes in amphibian populations, thereby improving the general understanding of genetics and mechanisms behind the evolution and maintenance of biological diversity. Phenotypic and genetic variation in life-history traits was studied within and between populations common frog (Rana temporaria) populations along a 1600 km transect from southern Sweden to northern Finland. Embryonic and larval development and growth was investigated both under field and laboratory conditions. The results suggest ample genetic diversity in larval life-history traits among Fennoscandian common frog populations. Larval developmental rate along the gradient has evolved a countergradient variation pattern of genotypes and phenotypes as indicated by the positive relationship between developmental rate and latitude under laboratory conditions and the lack of such a relationship in the field. The data suggest that this pattern has evolved because of time constraints due to decreasing length of growth season with latitude. Neither field-caught adults nor laboratory raised larvae displayed a linear latitudinal size cline as expected from the so called Bergmanns rule. Rather, size increased towards the mid-latitude populations and decreased thereafter, indicating that body size is a product of direct environmental induction or a trade-off with other life-history characters. Age and size at hatching showed no consistent latitudinal pattern, indicating that the embryonic stage is not as time constrained as the larval stage. A large part of the variation in age and size at metamorphosis among populations was due to additive genetic effects. However, small, but significant maternal effects, mostly due to variation in egg size and non-additive genetic effects also contributed to among population variation. A comparison of divergence in presumably neutral molecular genetic markers (FST) and quantitative characters (QST) revealed that although both estimates of divergence were relatively high, estimates of QST was generally higher than those of FST, indicating that the genetic variation observed in larval traits is primarily a result of natural selection rather than genetic drift. Hence, our results reinforce the conclusion that intraspecific genetic heterogeneity in the young northern European ecosystems may be more widespread than previously anticipated

Lichens are fascinating symbiotic systems, where a fungus and a unicellular alga, most often green (bipartite green algal lichens; 90% of all lichens), or a fi lamentous cyanobacterium (bipartite cyanobacterial lichens; 10% of all lichens) form a new entity (a thallus) appearing as a new and integrated organism: in about 500 lichens the fungus is associated with both a cyanobacterium and an alga (tripartite lichens). In the thallus, the lichen bionts function both as individual organisms, and as a symbiont partner. Hence, in lichens, the participating partners must both be able to receive and acquire resources from the other partner(s) in a controlled way. Lichens are particularly successful in harsh terrestrial environments. In part this is related to their poikilohydric nature and subsequent ability to repeatedly become desiccated and hydrated. Metabolic activity, i.e. photosynthesis, respiration, and for cyanobacterial lichens N2-fixation, is limited to periods when the thallus is suffi ciently hydrated. Mineral nutrients are mainly acquired from dry or wet deposition directly on the thallus. Taken together it then appears that lichens are to a large extent passively controlled by their environment, making their control over resource allocation and acquisition particularly challenging. The aim of this thesis was to investigate resource acquisition and allocation processes in different lichens, and to see how these respond to changes in resource availability. This was done by following lichen growth in the fi eld during manipulation of water, light, and nutrient supply, and by assessing the responses of both the integrated thallus as well as the individual bionts. As a fi rst step, resource allocation and acquisition was investigated for a broad range of lichens aiming to determine the magnitude of metabolic variation across lichens. Seventy-fi ve lichen species were selected to cover as broad a spectrum as possible regarding taxonomy, morphology, habitat, and nitrogen requirements. The lichens had invested their nitrogen resources so that photosynthetic capacity matched respiratory carbon demand around a similar equilibrium across the contrasting species. Regulation of lichen growth was investigated in another study, using the two tripartite species Nephroma arcticum and Peltigera aphthosa, emphasizing the contribution of both internal and external factors. The empirical growth models for the two lichens were similar, showing that weight gain is to a higher extent dependent on those external factors that regulate their photosynthesis, whilst area gain is more controlled by internal factors, such as their nitrogen metabolism. This might be inferred from another study of the same species, where nitrogen manipulations resulted in an undisturbed weight gain, a similar resource allocation pattern between the bionts, but a distorted area gain. Aiming to investigate lichen nitrogen relations even further, lichens’ capacities to assimilate combined nitrogen in the form of ammonium, nitrate and amino acids were assessed using 14 contrasting boreal species. All these had the capacity to assimilate all the three nitrogen forms, with ammonium absorption being more passive, and nitrate uptake being low in bipartite cyanobacterial lichens. Differences in uptake capacities between species were more correlated to photobiont than to morphology or substrate preferences. Finally, to investigate intra-specifi c plasticity in relation to altered nutrient supply, resource investments between photo- and mycobiont were investigated in the two bipartite green algal lichens Hypogymnia physodes and and Platismatia glauca in a low and a high nutrient environ- in a low and a high nutrient environ- ment. In both species, more of the resources had been directed to the photobiont in the high nutrient environment also increasing their overall carbon status. Taken together, my studies indicate that in spite of the apparent passive environmental control on lichen metabolism, these symbiotic organisms are able to both optimize and control their resource acquisition and allocation processes.

Multiple mating by females is common in nature. Yet, the evolution and maintenance of polyandry remains a bit of an evolutionary puzzle. It was my aim in this thesis to reach a greater understanding of this phenomenon as well as to investigate the consequences of polyandry on the evolution of reproductive divergence in insects. In an extensive meta analysis addressing the direct effects of multiple mating on female fitness in insects, I found that insects gain from multiple matings in terms of increased lifetime offspring production. In species without nuptial feeding, increased mating rate leads to decreased female lifespan and my results strongly support the existence of an intermediate optimal female mating rate. However, results from an experimental study where I examined the relationship between female fitness and mating rate in the bean weevil (Callosobruchus maculatus) showed that female fitness was maximized at two alternative mating rates, indicating that some species may exhibit a more complex relationship between the costs and benefits of mating. In the meta analysis on species with nuptial feeding, I found only positive effects of increased mating rate and the puzzle is rather what constrains the actual mating rates of females in these groups. Sexual selection is a very potent driver of rapid evolutionary change in reproductive characters. Most research has focussed on precopulatory sexual selection, but in promiscuous species sexual selection continues after copulation and variance in male fertilization success gives rise to postcopulatory sexual selection. In this thesis I found that three allopatric populations of the red flour beetle (Tribolium castaneum) have diverged in traits related to reproduction. Male genotype affected all aspects of female reproduction, but more interestingly, males and females interacted in their effect on offspring production and reproductive rate, showing that the divergence was due to the evolution of both male and female reproductive traits. When studying postcopulatory sexual selection, sperm competition has been put forward as the main source of variance in fertilization success. The results from a set of double-mating experiments, using the same populations of flour beetles, provided strong evidence that cryptic female choice is also important in generating variance in male fertilization success. I found not only main effects of female genotype on male fertilization success but also male-female interactions which provide more unambiguous evidence for cryptic female choice. Finally, I attempted to uncover which male signals-female receptors are involved in the reproductive divergence observed in the Tribolium populations. In a double-mating experiment I manipulated female perception of two male reproductive signals, copulatory courtship and cuticular hydrocarbons, and the results indicate that, within populations, both signals are sexually selected. However, only male cuticular hydrocarbons seem to be involved in the reproductive divergence between the populations. In conclusion, multiple mating by female insects can be understood solely in terms of direct fitness benefits resulting from increased offspring production. I have shown that postcopulatory sexual selection can lead to rapid divergence in reproductive traits related to mating and that cryptic female choice plays an important role in this divergence.

Temporal and spatial variations of pelagic microorganisms in the northern Baltic Sea were studied, as well as factors influencing their abundance and growth rates. Three main questions were asked 1) How does increased productivity influence the structure of the microbial food web? 2) Does predation limitation vary between trophic levels? 3) What is the relative importance of resource and predation limitation at different trophic levels? A field study in the northern Baltic Sea showed that dominating protozoa, flagellates and ciliates, increased with increasing primary productivity from north to south. Furthermore, relatively small protozoan cells dominated in the low productive north, while larger cells became more dominant in the south. The relationship between plankton size structure and productivity was further studied in an experimental system. In agreement with present theories regarding nutrient status of pelagic food webs, increased productivity caused a lengthening of the food chain as well as a change in plankton size structure. While microplankton dominated in nutrient rich treatments pico- and nanoplankton dominated during nutrient poor treament. The flagellate community was dominated by a potentially mixotroph, Chrysochromulina sp., at low nutrient concentrations. To our knowledge this is the first experimental study showing that Chrysochromulina sp. in resemblance with other mixotrophs is favoured by nutrient poor conditions compared to strict autotrophs and heterotrophs. During a stratified summer period autotrophic microorganisms in the northern Baltic Sea did not respond to removal of potential predators, indicating that they were primarily limited by inorganic nutrients. An exception was small eucaryotic picoplankton that showed a large response to predator removal. Among the heterotrophic microorganisms direct effect of predation seemed to increase from ciliates, heterotrophic bacteria, small heterotrophic flagellates, medium flagellates to large flagellates. No quick indirect effect was observed, but after four days trophic cascades were detected. The relative importance of resource and predation limitation was studied among heterotrophic bacteria, flagellates and ciliates in the northern Baltic Sea. For all these groups, resource limitation seemed to prevail during the summer period. The results also indicated that the relative importance of predation increased with the productivity of the system. To our knowledge there are no earlier measurements on the relative importance of resource and predation limitation for micoorganisms in the pelagic environment.

This thesis examines the relative importance of local conditions, dispersal and dynamics of the trees on epiphyte distribution patterns and colonization-extinction dynamics. Study species are the mosses Orthotrichum speciosum and O. obtusifolium, and the red-listed Neckera pennata. The thesis also includes an attempt to parameterize a model for a lichen metapopulation (Lobaria pulmonaria) in a dynamic landscape, based on only presence/absence data of the epiphyte and its host trees. The results show that epiphyte colonization of trees is affected by both local conditions, and by connectivity to occupied trees. The positive effect of connectivity, implying a restricted dispersal range, was established by both demographic and genetic studies. The important local conditions were tree diameter and vitality, and shade. Local extinctions from trees occurred among small trees with low local epiphyte abundance, but more often, were the results of tree fall. The observed importance of connectivity on epiphyte colonization agrees with the assumptions of the classic metapopulation model. However, the classic metapopulation model assumes that the landscape is static, and that local extinctions occur for stochastic reasons. The dynamics of epiphytes are different. A new conceptual model is therefore suggested, the patch-tracking metapopulation model. It differs from the classic metapopulation model in that it includes dynamics of the patches, and in that local extinctions only occur as patches are destroyed. Simulations of the dynamics of N. pennata showed that its future metapopulation size will be overestimated unless the dynamics of the trees are accounted for. The simulation results further suggest that the dynamics of N. pennata can be characterised by the patch-tracking metapopulation model. The attempt to parameterize the L. pulmonaria metapopulation model showed that more information are required for rigorous parameterization, preferably of the past historic fire regime.

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

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.