The social organization of animal societies has important implications for several fields of biology, from managing wild populations to developing new ecological and evolutionary theory. Although much attention has been given to the formation and maintenance of societies of group living individuals, less is known about how societies of solitary individuals have been shaped and maintained. Traditionally, the evolution of social organizations in the mammalian order Carnivora has been regarded as a directional selection process from a solitary ancestry into progressively more advanced forms of sociality. In this thesis, I tested this model against an alternative model, assuming radiation from a socially flexible ancestry. I further explored sociality, resource use and dispersal of a solitary carnivore, the wolverine (Gulo gulo), in the light of these two evolutionary models. Phylogenetic reconstruction generally supported that carnivore social organizations evolved through directional selection from a solitary ancestor. However, results from captive wolverine females indicated that they might have rudimentary social tendencies, which rather support that sociality in carnivores radiated from a socially flexible ancestry. Wild wolverines in northwestern Brooks Range, Alaska, adhered to the commonly found ecological niche as a largely ungulate dependent generalist carnivore. Lack of sexual asymmetry in dispersal tendencies indicated that resource competition among wolverine females probably was high. I suggest that wolverines have latent abilities to aggregate, but that their phylogenetic legacy in terms of morphology has constrained them into an ecological niche where resource abundance and distribution generally inhibit aggregations. Due to contradictory results, I suggest further research to test evolutionary theory regarding carnivore social evolution, and particularly to explore new avenues into social evolution that better explain intra-specific variation in sociality, as well as formation and maintenance of solitary social systems.

Natural stress and disturbances are important factors affecting the structure and function of ecosystems. However the magnitude of stress has escalated due to anthropogenic activities. Environmental monitoring and toxicity assessments try to protect ecosystems from unwanted human alterations. The aim of this Doctoral thesis was to increase the understanding of the complex effects that environmental stress has on individuals and invertebrate populations. The low saline environment in the Baltic Sea is perceived as stressful for most organisms living there. In Paper I, it was found that Baltic blue mussels living in the less saline northern Baltic Proper (~5 psu) had lower basal metabolism and were more susceptible to toxic exposure than the mussels in the south (~7 psu). There was no genetic differentiation between the mussels from the northern and southern areas while there were genetic differences between mussels from sites within the respective areas (Paper III), indicating that there is not a simple relationship between the health of the mussels and genetic diversity in the microsatellite loci studied. In Paper IV it was found that the heat tolerance of the intertidal dogwhelk Nucella lapillus is oxygen dependent. Increased oxygen levels resulted in higher survival rate. Protein expression profiles also became more similar to those of the controls, compared to the whelks exposed to high temperature and normal oxygen levels. In Paper V and VI it was found that exposure to a single toxicant for more than one generation decreased the genetic diversity in exposed copepod populations even though abundances remained unaltered. In Paper VI, exposure to naturally contaminated sediments, which contained of a mixture of toxicants, did not decrease genetic diversity. However the genetic divergence (FST) within the treatments was very high, probably due to small effective population sizes in the replicates. Likewise in Paper III, the very low blue mussel abundance in the north together with the stressful environment suggests a small effective population in the northern Baltic Proper. In conclusion, my studies show that, measuring effects on several levels, including both functional and structural endpoints will both increase the sensitivity of the tests and increase their ecological relevance.

In this thesis we have studied speciation in three butterfly genera Polygonia (Nymphalidae, Nymphalini), Pararge (Nymphalidae, Satyrinae) and Celastrina (Lycaenidae: Polyommatinae). In the first paper a dated phylogeny, based on molecular data, of Polygonia was constructed. We found strong conflict between the nDNA and mtDNA datasets. Possibly this can be explained by ancestral and recent hybridizations between contemporary taxa. The results point to the importance in using different markers when we try to resolve evolution of taxa. In the second paper a sister group comparison was made in order to discover whether host plant range has had an effect on species diversity in Polygonia. Our result indicated higher diversification rates in clades which included species with larvae feeding on different, or additional, plants compared to the ”urticalean rosids” specialists. In the third paper our focus was on the colonization abilities in polyphagous butterflies. The haplotype structures of the mtDNA cytochromeoxidase I (COI) within the Nearctic species of Celastrina as well as within P. c-album and P. faunus were analysed in a network. We found little variation in Celastrina and P. c-album. This results imply that the genera have expanded recently and rapidly. There are indications of differentiation in COI in Celastrina and, possibly, host plant use is involved. However, in P. faunus we found structure among the haplotypes. We believe that several different haplotypes of this species have been preserved during glaciations in the Nearctic. In the fourth paper the evolution of the grassfeeding Pararge was analysed. The phylogeny was based on the mtDNA COI and the nDNA wingless (wgl) and times of divergences were calculated. We found a deep divergence between the European and Moroccan populations of P. aegeria which indicates the importance of the Mediterranean as a barrier for gene flow.

The climate change is expected to affect especially alpine areas negatively, replacing the alpine flora with subalpine forest. The understanding of how vegetation influences total organic carbon (TOC) in soil, streams and lakes in alpine and subalpine areas will lead to a better understanding of the effects of climate change, and will also increase the knowledge of the ecotone as a whole. In this study plant-soil relations were examined in a subalpine and an alpine catchment in the north of Sweden, by comparing dissolved organic carbon (DOC) concentrations, 13C-DOC, 13 compared with lake and stream water DOC, as well as sediment OC from the recipient lakes in the catchments.  The results show that subalpine forests at lower altitudes, have higher DOC concentrations, higher C:N ratios, and more depleted Particulate OM in water and inlets, show that allochthonous carbon influences water properties in both catchments, as does primary production by benthic and pelagic algae, separating shallow and deep sediment Differences between the catchments are explained with the higher primary production of organic material and root exudations from trees in the subalpine forested catchment effecting the whole catchment dynamics. C-SOM and the carbon to nitrogen (C:N) ratios. The terrestrial bulk chemical properties of DOC were alsoδ13C signals in soil, and soil-solution compared to alpine areas. δ13C signals from Dissolved OM andδ13C signals.

Energiflöden mellan terrestra och akvatiska ekosystem har uppmärksammats mycket den senaste tiden. I den här studien undersöker jag betydelsen av ett terrestert djur, älgen (Alces alces), som dör i vattendrag och avger näring till vattnet. Genom litteraturstudier och teoretiska beräkningar uppskattade jag hur vanligt det är att älgar dör i vattendrag och om dessa älgar har någon inverkan på vattendragets totala näringsbudget. Jag fann att drunkning hos älgar tas upp som en parameter för naturliga dödsorsaker i flera artiklar från USA, Canada och Polen. Orsakerna till att älgar drunknar, trots att de är duktiga simmare, kan vara t.ex. att stranden är för brant, att de tvingas ner i vattnet av rovdjur, att skadeskjutna älgar söker sig till vatten på grund av törstkänningar i samband med blodförlust eller att de går igenom svaga isar. Utifrån teoretiska beräkningar fann jag även att en älg som dött i ett vattendrag bidrar till mycket liten del av den totala näringsbudgeten. Min hypotes är att det endast blir en lokal påverkan nedströms djuret. Hur denna påverkan ser ut och om den kommer primärproducenter och fiskar tillgodo är mycket komplext och undersöks inte i denna studie.

Statistical modelling is often used to relate the presence or abundance of species to environmental predictors, thereby providing a basis for predictive mapping of species or biodiversity. The variables included must thus be relevant and reflect actual changes in the environment. Therefore, the quantification of species–environment relationships is an important aspect of predictive modelling. This thesis examines how phytobenthic species or communities in the Baltic Sea relate to environmental gradients, and if different aspects of phytobenthic species distribution in the Baltic Sea could be explained by spatial or temporal variation in environmental factors. Predictive distribution modelling usually focuses on how environmental variables control the distribution of species or communities. Thus the relative weight of the predictor variables on different scales is of importance. In this thesis, I show that the relative importance of environmental variables depends both on geographic scale and location, and that it also differs between species or species groups. There are no simple explanations to the temporal variability in species occurrence. I here show that the temporal changes in species distribution within the phytobentic zone varies in a spatial context. I also try to find temporal and spatio-temporal patterns in species distribution that could be related to changes in climate or anthropogenic disturbance. However, the findings in this thesis suggest that single factor explanations are insufficient for explaining large-scale changes in species distribution. A greater understanding of the relationship between species and their environment will lead to the development of more sensitive models of species distributions. The predictions can be used to visualise spatial changes in the distribution of plant and animal communities over time. / At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Submitted. Paper 3: Manuscript. Paper 4: Manuscript. Paper 5: Manuscript.

The impacts of two strains of the dinoflagellate Alexandrium tamarense, differing in lytic activity, on the abundance and the composition of microbial communities (<150 μm) were studied in North Sea water during spring with Phaeocystis globosa as a dominant species. Cell-free suspensions (supernatant) of exponentially growing lytic and non-lytic Alexandrium culture were added at different concentrations to natural microbial communities under nutrient rich conditions. The non-lytic strain had a positive impact on diatoms whereas the lytic strain suppressed phytoplankton growth in comparison to the control. P. globosa, present as single cells in the initial community, increased in abundance and formed colonies in all treatments. However, total abundance and number of colonies was low with lytic Alexandriumadditions, whereas shape of the colonies, but not abundance of cells, was affected by non-lytic Alexandrium additions. During the 4-day experiment, bacterial abundance was constantly higher with high lytic additions (highest concentration equivalent to 1000 cells ml−1) whereas nanoflagellate abundance in the same treatments was found to be lower at the end of the experiment. Initial bacterial community composition differed significantly among lytic Alexandrium, non-lyticAlexandrium and North Sea water. However, neither bacterial activity nor composition was significantly affected by the supernatants after 96 h. Our results indicated that Alexandrium allelochemicals do not inhibit growth and production of bacteria in seawater collected during spring in the North Sea. / Allelopathy among phytoplankton - a structuring force among phytoplankton

A solid-phase extraction (SPE) method was developed for the determination of organochlorine pesticides, namely aldrin, alpha-BHC, beta-BHC, delta-BHC, dieldrin, endosulfan I, endosulfan II, endosulfan sulfate, endrin, endrin aldehyde, lindane, heptachlor, heptachlor epoxide, 4,4’-DDD, 4,4’-DDE and 4,4’- DDT in water. The effect of extraction conditions, such as the addition of sodium chloride and methanol to the sample prior to loading was studied. The sample was concentrated by a plain polystyrene-divinylbenzene resin, and the extract was eluted by ethyl acetate. Qualification and quantification of the target pesticides were performedby gas chromatography/ mass spectrometry (GC/MS) in the full-scan and selected ion-monitoring mode, respectively, and for better detection of pesticides in field samples the mass spectrometer was altered from electron ionization (EI) to chemical ionization mode (CI). The repeatability of the method for MilliQ-water fortified with pesticides at a level of 0.1 to 0.6 µg/l ranged from 8 to 18%, and the obtained recoveries ranged from 67 to 135%. The method was evaluated for the determination of organochlorine pesticides in fourteen surface- and groundwater samples taken from locations along King Talal Dam, King Abdullah Canal and Zarqa River in the Jordan Valley. The limit of detection of the pesticides residues in 500-ml field water samples ranged from 0.0009 to 15.7 ng/l. The obtained results confirmed the presence of trace amounts of some organochlorine pesticides in the analyzed samples, i.e. lindane and endosulfan compounds.

Våtmarker som anläggs på jordbruksmark kan förhindra att fosfor från åkrarna når sjöar och vattendrag och hjälper därmed till att minska övergödningen. Det är viktigt att utforma våt-marken på ett sådant sätt att så mycket som möjligt av den biotillgängliga fosforn hålls kvar. Syftet med studien var att undersöka om det finns ett positivt samband mellan lerinnehåll och fosfors biotillgänglighet i en anlagd våtmarks sediment. Syftet var även att undersöka om partikelstorleken minskar gradvis med avståndet från inloppet, det vill säga om mer lera sedimenterar närmare utloppet. Prover togs längs tre transekter från sedimentet i en våtmark i västra Östergötland och fosfor- samt partikelstorleksanalyser gjordes. Genom extraktion med ammoniumklorid (NH4Cl) kunde lättillgänglig fosfor bestämmas. Partikelstorleksanalysen gjordes med en modifierad pipettmetod, baserad på att större partiklar sedimenterar fortare än mindre partiklar. Regressionsanalys visade att partikelstorleken minskade signifikant från inlopp till utlopp i våtmarken. Däremot hittades inte några signifikanta samband mellan partikelstorlek och lättillgänglig fosfor eller totalfosfor, i motsats till resultat i andra studier. Halterna av både lera och olika fosforfraktioner i sedimentet var relativt låga och likartade vid olika provpunkter i våtmarken vilket kan ha påverkat resultaten.

Effects of fishery closure on perch was detected as female perch was on average 5 cm longer in closed areas compared to non-closed areas. This could be a result of larger individuals surviving as an effect of reduced fishing. No effect was detected on the reproduction rate of YOY (young of the year) perch. A zooplankton survey revealed less amounts of the most important plankton groups for YOY perch in areas with failed reproduction, though the difference was statistically insignificant. The distribution of YOY perch and YOY/adult stickleback differed completely in this study with no area where the two species are both present. It is sketchy to draw any conclusions from this data but it remains a field for further investigation. The reproduction problem is concentrated to the outer parts of the archipelago which confirms previous studies of the geographic extension.