The ongoing climate changes are predicted to accelerate fast in arctic regions with increases in both temperatures and precipitation. Although the duration of snow cover is generally expected to decrease in the future, snow depth may paradoxically increase in those areas where a large amount of the elevated precipitation will fall as snow. The annual distribution and duration of snow are important features in arctic ecosystems, influencing plant traits and species interactions in various ways. In this thesis, I investigated the effect of snow on plants and their interactions with herbivores by experimentally increasing the snow cover by snow fences in three different habitats along an environmental gradient in Abisko, northern Sweden. I found that the snow cover mattered for plant quality as food for herbivores and herbivore performance. An enhanced and prolonged snow cover increased the level of insect herbivory on dwarf birch leaves under field conditions. Autumnal moth larvae feeding on leaves that had experienced increased snow-lie grew faster and pupated earlier than larvae fed with leaves from control plots. These findings indicated that plants from snow-rich plots produced higher-quality food for herbivores. My studies showed that differences in snow-lie explained parts of the within-year spatial and seasonal variation in plant chemistry and patterns of herbivory in this arctic landscape. The relationship between leaf nitrogen concentration and plant phenology was consistent between treatments and habitats, indicating that snow per se, via a delayed phenology, was controlling the nitrogen concentration. The relationship between leaf age and level of herbivory was positive in the beginning of the growing season, but negative in the end of the growing season, indicating an increasing importance of plant palatability and a decreasing importance of exposure time in determining the level of herbivory throughout the growing season. The concentrations of phenolics varied among habitats, treatments and sampling occasions, suggesting that these plants were able to retain a mosaic of secondary chemical quality despite altered snow conditions. Furthermore, the nutrient limiting plant growth, according to N:P ratio thresholds, appeared to shift from nitrogen to phosphorus along the topographic gradient from snow-poor ridges to more snow-rich heathlands and fens. Snow addition had, however, no significant effect on other nutrient concentrations than nitrogen and no significant effect on the leaf N:P ratio, indicating that differences in snow cover could not explain the variation in plant nutrient concentrations among habitats. In a five-year study, I found opposing inter-annual effects of increased snow on plant chemistry. In contrast to earlier results, the effect of snow-lie on plant nitrogen concentration was predominantly negative. However, the effect of increased snow cover on the level of herbivory remained predominantly positive. The strong within-year relationship between snow-melt date (via plant phenology) and plant nitrogen concentration and level of herbivory could not predict inter-annual variation in the effect of snow manipulation. I did not find any conclusive evidence for a single factor causing the inter-annual opposing effect of snow addition, but the results indicated that interactions with summer and winter temperatures might be important. In conclusion, this thesis showed that climate-induced changes in snow conditions will have strong effects on plant traits and plant-herbivore interactions. However, alterations in snow cover do not influence all plant traits and the effect may vary in time and space.

No description available.

In host-parasitoid systems, there is a continuous coevolutionary arms race where each species imposes a strong selection pressure on the other. The host needs to develop defence strategies in order to escape parasitism and the parasitoid must evolve counter-defence strategies in order to overcome the host’s immune defence and successfully reproduce. This makes host-parasitoid systems excellent model systems for understanding evolutionary processes underlying host race formation and speciation. In order to gain a better understanding of the complexity of host-parasitoid interactions several aspects must be considered, such as search behaviour and host selection in the parasitoid, the development of immune responses in the host and counter-defence strategies in the parasitoid. In this thesis, I investigate interactions and coevolution in a natural host-parasitoid system, consisting of five species of Galerucella leaf beetles and three species of Asecodes parasitoids, by combining behavioural ecology with chemical ecology and immunology. In the studies performed, I found that pheromone production and responses in the beetles are connected to the phylogenetic relatedness between the Galerucella species (Paper I). I found no evidence that Asecodes exploits the adult pheromone to locate host larvae, but observed an ability in the parasitoids to distinguish a better host from a less suitable one based on larval odors (Paper II). The studies also revealed large differences in immune competence between the Galerucella species, which were linked to differences in hemocyte composition in the beetle larvae (Paper III, IV). Further, the results suggest that parasitism success in polyphagous Asecodes is strongly affected by former host species of the parasitoid (Paper IV). In conclusion, the results of this thesis suggest an on-going evolution in both parasitoid virulence and host immune responses in the Asecodes-Galerucella system. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 4: Manuscript.</p>

Anthropogenic disturbances are ubiquitous in coastal marine ecosystems. As such, more intensive monitoring efforts are necessary to conserve these valuable habitats. Bioindicators, organisms that predictably respond to changes in environmental variables, may be utilized in monitoring efforts to assess ecosystem functioning. To incorporate organisms into monitoring programs as bioindicators managers need to first understand the difference between the natural phenology of the focal organisms and their responses to different forms of anthropogenic disturbance. To determine if gammaridean amphipods could be used as indicators of changes in environmental quality in sub-tropical seagrass ecosystems, I conducted spatial and temporal surveys of amphipod communities in south Florida. Amphipod community structure varied significantly across sites and seasons. Variation in community structure was largely driven by macrophyte biomass, food availability, seasonally variable factors (epiphyte abundance, dissolved oxygen, salinity, and temperature), water-column nitrogen concentration, and factors related to freshwater input, including low Thalassia testudinum and high Halodule wrightii densities, and salinity. Amphipods are also susceptible to mechanical damage in seagrass habitats and could be used as indicators of ecological functioning of a region. A major source of mechanical damage in seagrass ecosystems is caused by boat propellers. I simulated propeller scars in continuous seagrass beds to investigate the effects of scarring on seagrass ecosystem functioning. Seagrasses located adjacent to propeller scars experienced a shift in the limiting resource from light to phosphorus. Amphipod community structure, however, was not impacted by scarring, but amphipod density was reduced in fragmented patches. To determine if plant-herbivore interactions were impacted by propeller scarring, we removed amphipods from half of the experimental plots and measured epiphyte biomass and community composition. Top-down control on epiphyte biomass or community composition by amphipods was not affected by fragmentation, despite reduced amphipod densities. My dissertation research demonstrates that amphipods could be incorporated into existing management programs in sub-tropical seagrass ecosystems as environmental indicators. Reduced amphipod densities in fragmented seagrass beds suggests that amphipods could also be used as ecological indicators, but more research is needed to determine the extent of the impacts of fragmentation on higher trophic levels.

About 6% of all angiosperms are dioecious. This separation of sexual function to male and female individuals, and the fundamentally different patterns of reproductive resource allocation that follows that separation, are thought to have important ecological and evolutionary consequences for plant enemy interactions.  I have studied whether intersexual differences in susceptibility to natural enemies can be explained by intersexual differences in resource allocation. In cases when sexual dimorphic traits form the target resource of a particular enemy I expected the enemy to select the best resource. The study system is the perennial dioecious herb, Silene dioica (Caryophyllaceae) and three of its specialist natural enemies, two insect herbivores the fly Delia criniventris (Anthomyiidae) and the twirler moth Caryocolum viscariella (Gelechiidae) and one systemic anther smut fungus Microbotryum violaceum. All three share the same food recourse, the floral stems, of their host plant. I studied the interaction on nine islands in a rising Bothinan archipelago over seven consecutive years. Both herbivores attacked female plants more than male plants (D. criniventris, 32.8% females, 30.7% males; C. viscariella, 4% females, 2% males). This attack pattern was consistent over years and islands and also correlated with a number of sexually dimorphic traits suggesting that females offer the better resource. Herbivore attack had no effect on plant survival but a significant effect on re-flowering the following year. Non-attacked females had an estimated mean re-flowering rate of 30.2%, and non-attacked males of 31%. Herbivore-attacked females had an estimated re-flowering rate of 46% compared with 38.4% for males. Females showed a stronger compensatory response to attack and tended to re-flower more often than males. Attack rates differed markedly in the different stages of primary succession. They were consistently higher in the youngest zone and decreased in parallel to progressing succession. This zonal pattern of decreasing attack rates correlated with several plant attributes, a decrease in plant size and nitrogen content, and an increased content of secondary compounds, but not to host plant density. We failed to come up with a simple explanation for the spatial structure with chronic high attack rates in the younger zones. However, the consistent patterns in attack rate suggest that a suite of abiotic and biotic factors interact and reinforce the strength and direction of selection. In general females were more frequently diseased by the anther smut Microbotryum violaceum than males with two exceptions. Disease frequencies were male biased on islands with low disease levels and in one of the seven study years. The change in disease frequencies from male to female bias confirm earlier studies suggesting that the relative contribution of the two components of infection risk, disease encounter and per contact infection probability can vary with population disease level. The change in the proportions of diseased males and females that was observed in one of the study years, followed a year of extreme weather conditions (prolonged drought). Both sexes showed a similar decline in flowering but diseased females decreased more than diseased males. This difference in response can be explained if considering that disease is more resource demanding in females than in males. Except for resources needed for mycelial growth and spore production, in females resources are also needed to restructure their sex expression and produce anthers. My study shows (i) that in dioecious species traits that are sexually dimorphic are of great importance for understanding the outcome of interactions with natural enemies, (ii) that the strengths and directions of enemy-host plant interactions are strongly shaped by both biotic and abiotic conditions.

In this thesis, I combined field, common-garden and greenhouse experiments to examine the ecological and evolutionary consequences of plant-herbivore interactions in the perennial herb Lythrum salicaria. More specifically I examined (1) whether resistance and tolerance to damage from herbivores vary with latitude and are positively related to the intensity of herbivory in natural populations, (2) whether effects of herbivory on plant fitness vary with latitude, (3) whether populations are locally adapted and whether herbivory influences the relative fitness of populations, and (4) whether the intensity and effects of insect herbivory on reproductive output vary locally along a disturbance gradient and are associated with differences in plant resistance. A common-garden and a greenhouse experiment demonstrated that plant resistance decreased whereas plant tolerance increased with latitude of origin among populations sampled along a latitudinal gradient in Sweden. Oviposition and feeding preference in the greenhouse and leaf damage in the common-garden experiment were negatively related to natural damage in the source populations. Experimental removal of insect herbivores in three populations sampled along the latitudinal gradient demonstrated that intensity of herbivory and its effects on plant fitness decreased towards the north. A reciprocal transplant experiment among the same three populations showed that herbivory affected the relative fitness of the three populations, but did not detect any evidence of local adaptation. Instead the southernmost population had the highest relative fitness at all three sites. A herbivore-removal experiment conducted in nine populations in an archipelago in northern Sweden demonstrated that insect herbivory strongly influenced among-population variation in reproductive output. However, variation in resistance was not related to differences in intensity of herbivory at this spatial scale. Taken together, the results demonstrate that resistance and tolerance to herbivory vary with latitude but in opposite directions, that intensity of herbivory is a major determinant of flowering and seed output, and that the strength of herbivore-mediated selection varies among populations in Lythrum salicaria. They further indicate that both physical disturbance regime and latitudinal variation in abiotic conditions may strongly influence the performance and abundance of perennial herbs because of their effects on interactions with specialized herbivores.

In this thesis, I combined observational studies with field and greenhouse experiments to examine selection on life history traits and variation in tolerance and resistance against herbivores in natural populations of the annual herb Arabidopsis thaliana in its native range. I investigated (1) phenotypic selection on flowering time and plant size, (2) the effects of timing of germination on plant fitness, (3) the effect of leaf damage on seed production, and (4) correlations between resistance against a specialist and a generalist insect herbivore. In all three study populations, flowering time was negatively related to plant fitness, but in only one of the populations, significant selection on flowering time was detected when controlling for size prior to the flowering season. The results show that correlations between flowering time and plant fecundity may be confounded by variation in plant size prior to the reproductive season. A field experiment detected conflicting selection on germination time: Early germination was associated with low seedling survival, but also with large leaf rosette before winter and high survival and fecundity among established plants. The results suggest that low survival among early germinating seeds is the main force opposing the evolution of earlier germination, and that the optimal timing of germination should vary in space and time as a function of the relative strength of selection acting during different life-history stages. Experimental leaf damage demonstrated that tolerance to damage was lowest among vegetative plants early in the season, and highest among flowering plants later in the season. Given similar damage levels, leaf herbivores feeding on plants before flowering should thus exert stronger selection on defence traits than those feeding on plants during flowering. Resistance against larval feeding by the specialist Plutella xylostella was negatively correlated with resistance against larval feeding by the generalist Mamestra brassicae and with resistance against oviposition by P. xylostella when variation in resistance was examined within and among two Swedish and two Italian A. thaliana populations. The results suggest that negative correlations between resistance against different herbivores and different life-history stages of herbivores may contribute to the maintenance of genetic variation in resistance.

Reindeer can have strong effects on the plant species composition and functioning of tundra ecosystems, and often promote a transition towards a graminoid-dominated vegetation type. As a result, they influence many ecological processes, such as nutrient dynamics, soil biotic composition and functioning, and carbon storage. Several studies suggest that the effect of reindeer on vegetation may follow predictable patterns and could induce an alternative stable vegetation state. However, little empirical data on the long-term stability of reindeer effects on vegetation exist, as it is inherently challenging to study these ecological processes experimentally on a sufficiently long timescale. The main objective of this thesis was therefore to gain a better understanding of the long-term ecological processes following reindeer-induced vegetation shifts. In order to gain a more mechanistic insight in what initially drives this transition, I used a field-based grazing simulation experiment in which I separated defoliation, trampling, moss removal and the addition of feces. This allowed me to test the relative contribution of reindeer-related activities to initiating the shift from moss and heath- dominated tundra towards a graminoid-dominated vegetation state. Additionally, I studied the long-term ecological stability following such a vegetation shift. I did this by addressing historical milking grounds (HMGs): sites where high reindeer concentrations associated with historical traditional reindeer herding practices induced a vegetation transition from shrubs towards graminoids several centuries earlier, but which were abandoned a century ago. Studying HMGs allowed me to address: 1. The potential stability of reindeer-induced vegetation shifts; 2. The ecological mechanisms contributing to the long-term stability of these vegetation shifts; and 3. How such long-lasting vegetation changes influence soil carbon- and nutrient cycling. I found that trampling by reindeer is an important mechanism by which reindeer cause vegetation change. Addressing HMGs further revealed that this vegetation change can be hightly persistent, as the studied HMGs showed only a low encroachment at the surrounding borders in the last 50 years. The vegetation in the core areas of all studied HMGs had remained strikingly stable, and were hardly invaded by surrounding shrubs. Interestingly, soil nutrient concentrations and microbial activities were still different from the surrounding area as well, and even comparable to actively grazed areas. Even after many centuries of changed vegetation composition and soil processes, there was no difference in total carbon sequestration. This suggests that the environmental conditions for microbial decomposition were more important than vegetation composition for the soil carbon stocks, in our study site. After studying the contemporary habitat use of HMGs by reindeer and other herbivores, investigating the potential plant-soil feedbacks mechanisms and detailed soil analyses, I concluded that several ecological mechanisms contribute to the long-term stability of HMGs: first, the altered soil biotic and abiotic conditions appear to have a stronger advantage for HMG vegetation than for the surrounding tundra vegetation. Furthermore, I found a clear browsing preference of small rodents on single shrubs proliferating in HMGs, causing a strong limitation on shrub expansion. Moreover, the dense established sward of graminoids likely poses a strong direct competition for space and nutrients, hindering seedling establishment. Finally, I conclude that HMGs are highly stable on relevant ecological timescales, and propose how the concepts of historical contingency and ASS can be applied to understand stability of these reindeer-induced vegetation transitions.

No description available.

No description available.