<p>Modern methods for managing pine (<em>Pinus sylvestris</em>) create homogenized forests. This decreases nature’s potential for biodiversity and might threaten species in need of different types of milieu. The main purpose of this study was to investigate how important older pine trees are for biodiversity. In the Hall-Hangvar Reserve in the north-west part of Gotland, insects collected from traps showed that more species were found in old or dead trees compared to younger pine trees. A statistically significant difference was found for Coleoptera (beetles). The taxons of greatest interest for this study were Coleoptera and Hymenoptera (wasps). Certain families of Hymenoptera use ducts made by larvae from some families of Coleoptera.These larvae also serve as prey. Relevance concerning enviromental importance to species and diffrenences in inhabiting the three stages of pine trees was of importance.</p>

Despite general concern that the massive deposits of methane stored under permafrost underground and undersea could be released into the atmosphere due to rising temperatures attributed to global climate change, little is known about the methanogenic microorganisms in permafrost sediments, their role in methane emissions, and their phylogeny. The aim of this thesis was to increase knowledge of uncultivated methanogenic microorganisms in submarine and terrestrial permafrost deposits, their community composition, the role they play with regard to methane emissions, and their phylogeny. It is assumed that methanogenic communities in warmer submarine permafrost may serve as a model to anticipate the response of methanogenic communities in colder terrestrial permafrost to rising temperatures. The compositions of methanogenic communities were examined in terrestrial and submarine permafrost sediment samples. The submarine permafrost studied in this research was 10°C warmer than the terrestrial permafrost. By polymerase chain reaction (PCR), DNA was extracted from each of the samples and analyzed by molecular microbiological methods such as PCR-DGGE, RT-PCR, and cloning. Furthermore, these samples were used for in vitro experiment and FISH. The submarine permafrost analysis of the isotope composition of CH4 suggested a relationship between methane content and in situ active methanogenesis. Furthermore, active methanogenesis was proven using 13C-isotope measurements of methane in submarine permafrost sediment with a high TOC value and a high methane concentration. In the molecular-microbiological studies uncultivated lines of Methanosarcina, Methanomicrobiales, Methanobacteriacea and the Groups 1.3 and Marine Benthic from Crenarchaeota were found in all submarine and terrestrial permafrost samples. Methanosarcina was the dominant group of the Archaea in all submarine and terrestrial permafrost samples. The archaeal community composition, in particular, the methanogenic community composition showed diversity with changes in temperatures. Furthermore, cell count of methanogens in submarine permafrost was 10 times higher than in terrestrial permafrost. In vitro experiments showed that methanogens adapt quickly and well to higher temperatures. If temperatures rise due to climate change, an increase in methanogenic activity can be expected as long as organic material is sufficiently available and qualitatively adequate. / Trotz allgemeiner Bedenken, dass auf Grund des Temperaturanstieges im Zusammenhang mit der globalen Klimaerwärmung große Mengen des in terrestrischen und submarinen Permafrostsedimenten gespeicherten Methans freigesetzt werden könnte, ist bisher wenig über die in diesen Böden lebenden methanogenen Mikroorganismen, ihre Phylogenese und sowie ihre Bedeutung hinsichtlich der Methanemissionen bekannt. Das Ziel dieser Doktorarbeit war die Erweiterung der bisherigen Kenntnisse über unkultivierte methanogene Mikroorganismen in submarinen und terrestrischen Sedimentablagerungen, die Zusammensetzung ihrer Lebensgemeinschaft, ihrer Phylogenese und ihrer Bedeutung bei der Emission von Methan. Es wird vermutet, dass methanogene Gemeinschaften submarines Permafrostes zur Erstellung von Modellen genutzt werden können, um Aussagen bezüglich potenzieller Reaktionen methanogener Gemeinschaften des kälteren terrestrischen Permafrostes auf steigende Temperaturen, zu ermöglichen. Die Zusammensetzung der methanogenen Gemeinschaft wurde in terrestrischen und submarinen Permafrostproben untersucht. Der im Rahmen dieser Forschungsarbeit untersuchte submarine Permafrost wies eine im Vergleich zum terrestrischen Permafrost um circa 10°C höhere Temperatur auf. Mittels Polymerasenkettenreaktion (PCR) wurde von jeder der Proben DNA extrahiert und mittels weiterer molekular-mikrobiologischen Methoden wie DGGE, RT-PCR und Klonierung analysiert. Des Weiteren wurden die Proben für in vitro Experimente und Zellzählungen (DAPI und FISH) verwendet. Die Analyse der Isotopenzusammensetzung von CH4 in submarinen Permafrostsedimenten ließ einen Zusammenhang zwischen Methangehalt und aktiver in situ Methanogenese vermuten. Überdies konnte aktive Methanogenese, mittels 13C-Isotopenmessungen von Methan in submarinem Permafrostsediment mit hohem TOC-Wert und hoher Methankonzentration, bewiesen werden. Im Rahmen der molekular-mikrobiologischen Untersuchungen wurden in allen submarinen und terrestrischen Permafrostproben unkultivierte Linien von Methanosarcina, Methanomicrobiales, Methanobacteriacea und den Gruppen 1.3 und Marine Benthic von Crenarchaeota gefunden. Methanosarcina war in allen submarinen und terrestrischen Permafrostproben die dominierende Gruppe der Archaeen. Die Zusammensetzung der archaealen Gemeinschaft, insbesondere die Zusammensetzung der methanogenen Gemeinschaft, variierte zwischen den submarinen und terrestrischen Proben. Des Weiteren fand sich bei der Zellzählung der Methanogenen im submarinen Permafrost eine 10-fach höhere Zellzahl als im terrestrischen Permafrost. Die in vitro Experimente zeigten, dass Methanogene sich schnell und gut an höhere Temperaturen anpassen können. Im Falle weiter steigender Temperaturen auf Grund der Klimaveränderungen, kann – bei ausreichender Verfügbarkeit und Qualität organischen Materials – mit einer Zunahme der methanogenen Aktivität gerechnet werden.

Lichens can colonize nearly all terrestrial habitats on earth and are functionally important in many ecosystems. Being poikilohydric, their active growth periods are restricted to periods when the thallus is hydrated from atmospheric water sources, such as rain, fog and high relative humidity. Since lichen hydration varies greatly over time lichen growth is therefore more difficult to model compared with, for example vascular plants with more even water supply. I developed two models to predict lichen hydration under field conditions that incorporates the atmospheric water potential (Ψair), derived from air temperature and humidity, only or in combination with species-specific rehydration and desiccation rates. Using Ψair allows the prediction of hydration induced by several water sources. These models were very accurate for epiphytic lichens with a close coupling to atmospheric conditions, but they were less accurate for mat-forming lichens with substantial aerodynamic boundary layers. The hydration model was further developed to include photosynthetic activation for different species, in order to compare their performance under different micro-climatic scenarios. Water balance and activation rate had large effects on lichen activity and were positively related to habitats providing long hydration periods, for example close to streams. To study effects of climate change, a complete model for net carbon gain (photosynthesis minus respiratory losses) was developed for an epiphytic lichen with intricate responses to light, hydration and temperature. Simulation responses in different climate scenarios revealed that projected climate change on a regional scale resulted in varied local scale responses. At the lighter, exposed sites of a forest, the growth responses were positive, but were potentially negative at darker sites with closed canopy. At the local scale, fluctuating hydration, summed irradiance when wet and Chlorophyll a are variables that predict lichen growth. However, at a landscape scale, these variables may be too detailed. We tested this for two terrestrial, mat-forming lichens and developed statistical models for lichen growth in the widest possible climatic gradient in northern Scandinavia, varying in light, temperature and precipitation. Light was the most important factor for high growth at the landscape scale, reaching saturation at a site openness of 40 %, equivalent to a basal tree area of 15 m2 ha -1 in this study. Thereafter, hydration was the next limiting factor, which could be well described by precipitation for one of the species. The simplest predictor was the normal temperature in July, which was negatively correlated with growth. It was apparent that the predictive variables and their power varied at different scales. However, light and hydration are limiting at all scales, particularly by light conditions when lichens are wet. This implies that ensuring that there is sufficient light below the forest canopy is crucial for lichen growth, especially for mat-forming lichens. Hydrophilic lichens may be better preserved in open habitats with long hydration periods. It was shown that models can be powerful and “easy to use” tools to predict lichen responses in various habitats and under different climate scenarios. Models can therefore help to identify suitable habitats with optimal growth conditions, which is very important for the conservation and management of lichens and their habitats. / Lavar kan kolonisera nästan alla terrestriska habitat i världen och är funktionellt viktiga i många ekosystem. Eftersom lavar är poikilohydriska (växelblöta), är deras aktiva tillväxtperioder begränsade till den tid då bålen är blöt från atmosfäriska vattenkällor, såsom regn, dimma och hög relativ fuktighet. Eftersom lavars vatteninnehåll varierar stort över tid är lavars tillväxt svårare att modellera jämfört med till exempel kärlväxter, med en mer jämn vattentillgång. Jag har utvecklat två fuktmodeller som förutsäger lavars vatteninnehåll i fält. Modellerna använder den atmosfäriska vattenpotentialen (Ψair), som erhålls från lufttemperatur och -fuktighet, antingen enbart eller i kombination med de artspecifika uppblötnings - och uttorkningshastigheterna. Genom att använda (Ψair) kan man förutsäga lavars vatteninnehåll från flera vattenkällor. Dessa modeller var mycket precisa för epifytiska lavar med en nära koppling till de atmosfäriska förhållandena, men fungerade mindre väl för mattlevande lavar med ett betydande gränsskikt. Fuktmodellen utvecklades ytterligare för att inkludera även fotosyntetisk aktivering av olika lavar, för att kunna jämföra deras aktivitet i olika mikroklimatiska scenarior. Vattenbalans och aktiveringshastighet hade stor effekt på på lavars aktivitet och var positivt relaterad till habitat med tillräckligt långa fuktperioder, till exempel habitat nära strömmande vatten. För att studera klimateffekter på lavar, utvecklade jag en total modell för nettoförvärv av kol (fotosyntes minus respiration) för en epifytisk lav med dess intrikata förhållande mellan ljus, fukt och temperatur. Simuleringar av modellen visade att lavens responser i förhållande till regionala klimatförändringar var kontrasterande på lokal nivå. Vid ljusa, öppna lokaler i skogen ökade tillväxten medan de potentiellt minskade vid mörka lokaler med ett mer slutet krontäcke. På den lokala skalan kan fluktuerande vatteninnehåll, summerat ljus när laven är blöt, och klorofyll a- innehåll förutsäga lavars tillväxt. Men, på en landskapsskala kan dessa variabler vara för detaljerade. Vi testade detta för två terrestriska, mattlevande lavar och utvecklade en statistisk modell för lavars tillväxt i en så stor klimatgradient som möjligt i norra Skandinavien genom att variera ljus, temperatur och nederbörd. Ljus var den viktigaste faktorn för att nå hög tillväxt på landskapsnivå där en mättnad nåddes vid 40 % öppenhet i skogen, som motsvarade en grundyta på 15 m2 ha -1 i den här studien. Fuktigheten var den näst viktigaste begränsande faktorn och kunde beskrivas väl med nederbörd för en av arterna. Den mest lättanvända faktorn var normaltemperaturen för juli månad, som i sin tur var negativt korrelerad till tillväxt. Det var tydligt att de prediktiva variablerna och deras förutsägande förmåga varierade med olika skalor. Ljus och fukt var begränsande på alla nivåer, speciellt av ljusförhållandena då lavarna är blöta. Detta innebär att tillräckligt höga ljusnivåer under krontäcket är avgörande för lavars tillväxt, speciellt mattlevande lavar. Hydrofila lavar torde bevaras bättre i öppna habitat med tillräckligt långa fuktperioder. Det var tydligt att modeller kan vara betydelsefulla och lättanvända verktyg för att förutsäga lavars responser i en bredd av habitat med olika mikroklimat. Modeller kan därför vara en hjälp för att identifiera lämpliga habitat med optimala tillväxtförhållanden och detta är viktigt för att bevara och sköta lavar och deras habitat.

Riparian vegetation along streams and rivers affects the aquatic community in numerous ways and often operates as a link for energy flux between forest and streams. The studies presented in this licentiate thesis focus on light and terrestrial invertebrates, two factors influenced by riparian zone structure, which potentially affect stream ecosystems and thus also brown trout (Salmo trutta). Paper I is a laboratory experiment where I study size dependent foraging behavior on surface-drifting terrestrial invertebrates and benthic invertebrates by brown trout. The results show a size-dependent difference in foraging ability with large trout being better able to use terrestrial surface prey than small trout. I argue that such ontogenetic foraging differences are due to both morphological constraints (eg. gape limitation) and size dependent behavioral differences related to predation risk. Paper II consists of a 5 month-long 2x2 factorial design field experiment where my objective was to examine the effects of terrestrial invertebrate input and solar radiation (PAR) on different trophic levels in a boreal headwater stream. More specifically, I followed the effects of increased light and decreased terrestrial invertebrate subsidies on periphyton, benthic macroinvertebrates and two size classes of the top fish predator, brown trout. The results showed that the reduction of terrestrial invertebrate input had size- and seasonal-dependent effects on trout, where large trout had lower growth rates than small trout, mainly in summer. Diet analyses of trout supported growth differences in that large trout in unmanipulated enclosures consumed relatively more terrestrial prey than large trout living in enclosures with reduced terrestrial inputs. A higher reliance on terrestrial prey subsidies by large trout compared to small may be explained by ontogenetic differences in foraging and habitat choice. Despite a 2.5-fold increase in PAR, light did not have an effect on chlorophyll a biomass, nor was there an effect on the density or composition of benthic macroinvertebrates. The lack of effects on primary production may be explained by very low nutrient levels in the stream.

Virtually all diurnal birds have tetrachomatic vision based on four different colour receptors. As a result, birds are potentially able to perceive their environment in twice as many colours as humans and four times as many colours compared to most other mammals, which are dichromatic. In addition to the spectrum visible to humans, birds are able to detect ultraviolet (UV) light. Signals with a UV component have been shown to be important to birds both in foraging and colour signalling. Because of the superior colour discrimination of the avian eye, UV sensitivity, but especially owing to its tetrachromacy, we cannot know what birds look like to those that matter, i.e. other birds. In my thesis I describe a new molecular method with which it is possible to identify the vision system of birds only using a small amount of DNA, without the need to keep or sacrifice the animal. It thereby facilitates large screenings, including rare and endangered species. The method has been used to increase the number of species with identified vision system type from 19 to 66. I show that raptors and songbirds have different vision systems, giving songbirds the possibility of a secret channel for colour signalling, and that male songbirds in coniferous forest take advantage of this to be significantly more cryptic to raptors than to females songbirds. I show that gulls have gained a vision system enabling them to detect the UV signals of fish when the fish swim close to the surface. Even though we tend to be rather self-satisfied with the quality of our colour vision, we are colour-blind when compared to birds. My work shows that human colour vision is inadequate for judging animal coloration, and that there is much more going on in bird colour signalling than meets our eye.

A low content of organic matter, which is largely refractory in nature, is characteristic of most sediments, meaning that aquatic deposit-feeders live on a very poor food source. The food is derived mainly from sedimenting phytodetritus, and in temperate waters like the Baltic Sea, from seasonal phytoplankton blooms. Deposit-feeders are either bulk-feeders, or selective feeders, which preferentially ingest the more organic-rich particles in the sediment, including phytodetritus, microbes and meiofauna. The soft-bottom benthos of the Baltic Sea has low species biodiversity and is dominated by a few macrobenthic species, among which the most numerous are the two deposit-feeding amphipods Monoporeia affinis and Pontoporeia femorata, and the bivalve Macoma balthica. This thesis is based on laboratory experiments on the feeding of these three species, and on the priapulid Halicryptus spinulosus. Feeding by benthic animals is often difficult to observe, but can be effectively studied by the use of tracers. Here we used the radioactive isotope 14C to label food items and to trace the organic matter uptake in the animals, while the stable isotopes 13C and 15N were used to follow feeding on aged organic matter in the sediment. The abundance of M. balthica and the amphipods tends to be negatively correlated, i.e., fewer bivalves are found at sites with dense populations of amphipods, with the known explanation that newly settled M. balthica spat are killed by the amphipods. Whether the postlarvae are just accidentally killed, or also ingested after being killed was tested by labelling the postlarvae with 14C and Rhodamine B. Both tracer techniques gave similar evidence for predation on and ingestion of postlarval bivalves. We calculated that this predation was likely to supply less than one percent of the daily carbon requirement for M. affinis, but might nevertheless be an important factor limiting recruitment of M. balthica. The two amphipods M. affinis and P. femorata are partly vertically segregated in the sediment, but whether they also feed at different depths was unknown. By adding fresh 14C-labelled algae either on the sediment surface or mixed into the sediment, we were able to distinguish surface from subsurface feeding. We found M. affinis and P. femorata to be surface and subsurface deposit-feeders, respectively. Whether the amphipods also feed on old organic matter, was studied by adding fresh 14C-labelled algae on the sediment surface, and using aged, one-year-old 13C- and 15N-labelled sediment as deep sediment. Ingestion of old organic matter, traced by the stable isotopes, differed between the two species, with a higher uptake for P. femorata, suggesting that P. femorata utilises the older, deeper-buried organic matter to a greater extent. Feeding studies with juveniles of both M. affinis and P. femorata had not been done previously. In an experiment with the same procedure and treatments as for the adults, juveniles of both amphipod species were found to have similar feeding strategies. They fed on both fresh and old sediment, with no partitioning of food resources, making them likely to be competitors for the same food resource. Oxygen deficiency has become more wide-spread in the Baltic Sea proper in the last half-century, and upwards of 70 000km2 are now devoid of macrofauna, even though part of that area does not have oxygen concentrations low enough to directly kill the macrofauna. We made week-long experiments on the rate of feeding on 14C-labelled diatoms spread on the sediment surface in different oxygen concentrations for both the amphipod species, M. balthica and H. spinulosus. The amphipods were the most sensitive to oxygen deficiency and showed reduced feeding and lower survival at low oxygen concentrations. M. balthica showed reduced feeding at the lowest oxygen concentration, but no mortality increase. The survival of H. spinulosus was unaffected, but it did not feed, showing that it is not a surface deposit-feeder. We conclude that low oxygen concentrations that are not directly lethal, but reduce food intake, may lead to starvation and death in the longer term.

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.

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.

No description available.