<p>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.</p><p>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.</p><p>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 <i>Nephroma arcticum</i> and <i>Peltigera aphthosa</i>, 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. </p><p>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 <i>Hypogymnia physodes </i>and and <i>Platismatia glauca</i> 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.</p>

<p>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.</p><p>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.</p><p>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. </p><p>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.</p>

<p>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?</p><p>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, <i>Chrysochromulina</i> sp., at low nutrient concentrations. To our knowledge this is the first experimental study showing that <i>Chrysochromulina</i> sp. in resemblance with other mixotrophs is favoured by nutrient poor conditions compared to strict autotrophs and heterotrophs.</p><p>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.</p><p>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.</p>

<p>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 </p><p>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. <i>Daphnia pulex</i> feeding on phytoplankton, and <i>Chydorus sphaericus</i> 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 <i>D. pulex</i>, the inflow of phytoplankton was made accessible to periphyton, and indirectly to <i>C. sphaericus</i>, which increased to such high densities that <i>D. pulex</i> 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.</p>

<p>Mängden död ved i brukade skogar har minskat med ungefär 90 %jämfört med naturliga skogar. Vid slutavverkning minskar mängden ytterligare. Att lämna naturhänsyn vid avverkning är ett viktigt steg för att bevara biologisk mångfald i skogslandskapet. Denna studie undersöker om naturhänsyn tas bort efter slutavverkning. På 63 % av de inventerade lokalerna hade naturhänsyn tagits bort, mestadels vindfälld tall. Det medförde en genomsnittlig minskning av mängden död ved med 2,0 m3 ha-1, vilket är en tredjedel av den totala volymen död ved lämnad vid slutavverkning. En enkät skickades till privata skogsägare. Den främsta orsaken till att man tagit bort träd var relaterade till ekonomi och produktion. Hur ekonomiskt beroende skogsägaren var av sin skog eller om man certifierat sin skog eller inte hade dock ingen betydelse för hur man hanterade naturhänsynen efter slutavverkning. Det berodde snarare på skogsägarnas attityd till produktion kontra bevarandet av biologisk mångfald. Skogsägare som tagit bort hänsynsträd var överlag mer tveksamma till att lämna olika former av naturhänsyn vid slutavverkning. Denna studie utvärderar även mängd och kvalitet på naturhänsyn lämnad efter slutavverkning i Östergötland.</p>

<p>Nutrient enrichment is the main cause of the increasing eutrophication process in the Baltic Sea during last century. A prerequisite counteract this process the load of nutrients to the Sea must be investigated. The large contribution of nutrients to the Sea is caused by emission from the rivers within the drainage basin. For the total emission of nutrient into the Sea the retention process in the rivers has big influence. This process in the river system has been assumed as a not important part of the nutrient cycle. However some investigators have pointed out that the retention process occurs in the river and be an important part of the nutrient dynamics in the river. For the object of investigation the Odra River Basin was chosen, as an example of a large river basin in the Baltic Sea Drainage Basin. The Odra River is mainly localized in Poland and contributes substantially to the eutrophication of the Baltic Proper. The Odra River is one of the least dammed major rivers of the Baltic Proper and as such of especial interest in retention studies. The phosphorus was chosen as an example of nutrient in the eutrophication process. The purpose of the investigation is to determine the phosphorus retention within the river. Two different models were used to calculate the retention process in the river. The"Multiple Regression Model"turned out to be insignificant, therefore the results were computed by using a “similarity model”, based on similarities between load from monitored tributary basins and the rest of the sub-basin. The retention in the Odra River in the amounted to -4608 t yr-1, which is 43% of the total phosphorus emission to the Odra River. There is also stated that in the Notec and the Warta rivers the retention process occur in the amount of -1940 t yr-1 and -3007 t yr-1 respectively.</p>

<p>Sedan 1992 har Jönköpings kommun drivit en naturskola i form av en exkursionsbuss, ekobussen. Verksamheten har varit inriktad mot dagsexkursioner för grundskolans senare år och gymnasieskolan under ledning av utbildad och engagerad personal. Genom åren har, i dialog med kommunens undervisande lärare, en pedagogisk modell för praktisk ekologiundervisning etablerats, där jämförande undersökningar av olika biotoper har utgjort tyngdpunkten. Man har använt ett undersökande och elevaktivt arbetssätt. Bussens utrustning har kunnat användas för avancerade mätningar och examination av djur och växter men tillhandahåller även personlig utrustning som stövlar och regnkläder. Det innebär stor flexibilitet i verksamheten. </p><p>I föreliggande studie berörs hur ett upplevelsebaserat lärande kan främja ett ökat miljömedvetande. I sex gymnasieklasser har undersökts i vilken mån uppfattningarna om den pedagogiska modellen förenas hos deltagarna. Vidare har undersökts hur exkursionen integreras i övrig undervisning i ekologi- och miljöundervisningen på skolorna. Slutligen besvaras frågan hur detta uppfattats av eleverna. Studien har både kvantitativa inslag i form av enkäter till cirka deltagande 150 elever, före och efter exkursion, och kvalitativa delar som innefattar deltagande observation under sex exkursioner, intervjuer med de sju lärare och även med initiativtagaren och den ansvarige för ekobussen. Undersökningen har huvudsakligen genomförts under maj månad år 2004. </p><p>Resultaten visar att verksamheten är mycket efterfrågad, både av lärare och av elever. Eleverna är i stort väl förberedda både teoretiskt och praktiskt och att upplevelserna under dagen används i efterarbetet. Hårt bokningstryck ger dock praktiska problem att inordna exkursionen i ordinarie undervisning. Exkursionsledaren kör bussen men styr också den pedagogiska verksamheten. Det finns en dynamik i processen att välja arbetsformer och parera för händelser under dagen. Det har över tid skett en förskjutning av fokus från kvantifiering av data till mera upplevelsebaserade erfarenheter. En tendens till passiv elevroll finnsi vissa situationer, när klassen lyssnar till exkursionsledarens genomgångar. Under praktiska arbetspass sker redovisningar genom att gruppen samtalar om resultaten, även om de flesta elever också får skriva en formell rapport i efterhand. </p><p>Det finns en stark ambition att vara en utomhuspedagogisk modell för skolorna. Man förefaller också lyckas vara den impuls till pedagogisk utveckling av arbetsformerna som styrdokumenten för skolan uppmuntrar till. Genom sin breda kontakt med kommunens lärare påverkar ekobussens pedagogiska modell ekologi- och miljöundervisningen på ett positivt sätt. Detta kan därmed vara ett värdefullt bidrag till ett ökat miljömedvetande.</p>

<p>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. </p><p>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. </p><p>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. </p><p>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.</p> / <p>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.</p><p>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.</p><p>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.</p><p>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.</p><p>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.</p>

<p>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.</p><p>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.</p><p>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.</p><p>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.</p><p>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.</p>

<p>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. </p><p>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.</p><p>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.</p><p>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.</p><p>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.</p>