Climate change is expected to have a strong impact on subarctic ecosystems. Increased temperatures as well as altered precipitation and snow cover patterns are predicted to change species distribution and affect biogeochemical processes in the subarctic tundra. Bryophytes are an essential vegetational component in northern ecosystems, due to their high abundance and importance in many ecological processes. In this study the effects of elevation and altered snow cover on the temporal dynamics of freezing damage in three subarctic bryophyte species (Hylocomium splendens, Ptilidium ciliare, and Sphagnum fuscum) were studied in a snow manipulation field experiment in Abisko, during early spring. Soil temperature and field moisture of moss shoots were collected. A freeze-thaw incubation experiment was conducted to investigate the freeze-thaw cycle resistance of H. splendens and P. ciliare originating from habitats with two differing snow-cover thicknesses. Freezing damage differed significantly between the bryophytes species with P. ciliare experiencing the least and S. fuscum the highest damage. Damage was higher at the low elevation, possibly attributable to acclimation effects. Snow removal led to higher damage in moss shoots, but no interactions of the different snow cover treatments with elevation, species or time were found. Freezing damage increased over time and no recovery occurred, likely due to temporal patterns in soil freeze-thaw cycles during early spring. Soil freeze-thaw cycles were the main factor influencing damage in bryophytes after snow melt. Measured environmental parameters could not explain the entire variation in damage. Damage might additionally be attributable to increased UV radiation or disturbances by herbivores.

This thesis examines the occurrence of hydrologic variability in subarctic ponds within the Churchill region of the Hudson Bay Lowland (HBL) and investigates the utility of using remote sensing studies to characterize changes in pond surface area. The thesis also characterizes hydro-climatic change over the past ~60 years, and compares this to pond sustainability within the region of Churchill. A multiple-methods approach incorporating field research, simple water balance modeling and remote sensing is used to address these objectives. Research findings demonstrate the occurrence of natural fluctuations in pond surface area and water levels in the Canadian subarctic. These fluctuations in pond water levels (and thus surface area) are caused by differences in antecedent hydrologic conditions, which are easily detected using remotely sensed imagery and may produce unrepresentative estimates of pond surface area change. Resulting from a 4.5 - 11.8 cm variation in water depth, pond surface areas were significantly altered by antecedent precipitation (average: 3,711 m²), intra-seasonal variability (average: 2,049 m²) and inter-annual climatic variations (average: 1,977 m²). These noteworthy pond boundary and water level differences reinforce the importance of accounting for hydrologic variability when delineating representative pond coverage and sustainability. Contemporary pond sustainability findings reveal significant regional climatic change, changing pond hydrologic conditions and overall pond physical stability between 1947 and 2008. Specifically, the Churchill region has become warmer and wetter. Occurring at a rate of 1.37 mm/yr over the study period, changing atmospheric conditions caused a decrease in open water pond hydrologic deficits. During the hydrologic recharge period, modeled pond water levels exhibited an increasing trend (August +0.72 mm/yr, September 0.51 mm/yr), which suggests ponds are filling closer to their maximum storage capacity prior to freeze-up. A remote sensing analysis of pond boundary modifications in mid-summer revealed no change in contemporary physical pond sustainability. Detected surface area changes from imagery were mainly attributed to naturally induced hydrologic variability. Overall, this thesis suggests a new methodological approach for conducting remote sensing pond sustainability research within the arctic/subarctic environment. As well, this study determined pond sustainability within the Churchill region over the last ~60 years.

This thesis examines the occurrence of hydrologic variability in subarctic ponds within the Churchill region of the Hudson Bay Lowland (HBL) and investigates the utility of using remote sensing studies to characterize changes in pond surface area. The thesis also characterizes hydro-climatic change over the past ~60 years, and compares this to pond sustainability within the region of Churchill. A multiple-methods approach incorporating field research, simple water balance modeling and remote sensing is used to address these objectives. Research findings demonstrate the occurrence of natural fluctuations in pond surface area and water levels in the Canadian subarctic. These fluctuations in pond water levels (and thus surface area) are caused by differences in antecedent hydrologic conditions, which are easily detected using remotely sensed imagery and may produce unrepresentative estimates of pond surface area change. Resulting from a 4.5 - 11.8 cm variation in water depth, pond surface areas were significantly altered by antecedent precipitation (average: 3,711 m²), intra-seasonal variability (average: 2,049 m²) and inter-annual climatic variations (average: 1,977 m²). These noteworthy pond boundary and water level differences reinforce the importance of accounting for hydrologic variability when delineating representative pond coverage and sustainability. Contemporary pond sustainability findings reveal significant regional climatic change, changing pond hydrologic conditions and overall pond physical stability between 1947 and 2008. Specifically, the Churchill region has become warmer and wetter. Occurring at a rate of 1.37 mm/yr over the study period, changing atmospheric conditions caused a decrease in open water pond hydrologic deficits. During the hydrologic recharge period, modeled pond water levels exhibited an increasing trend (August +0.72 mm/yr, September 0.51 mm/yr), which suggests ponds are filling closer to their maximum storage capacity prior to freeze-up. A remote sensing analysis of pond boundary modifications in mid-summer revealed no change in contemporary physical pond sustainability. Detected surface area changes from imagery were mainly attributed to naturally induced hydrologic variability. Overall, this thesis suggests a new methodological approach for conducting remote sensing pond sustainability research within the arctic/subarctic environment. As well, this study determined pond sustainability within the Churchill region over the last ~60 years.

<p> Evaporation was calculated for a subarctic beach ridge, near Churchill, Manitoba, using the energy balance approach. Energy balance calculations for the measurement season revealed an average Bowen ratio, β, of 0.68, with a value of 1.00 representing α' (the evaporability parameter). Fifty-seven percent of the net radiation was utilized by the evaporative heat flux over this tundra surface. Regressions were used to determine the most likely combination of environmental variables responsible for the behaviour of evaporation. Surface soil moisture remained relatively constant throughout the summer measurement period and soil temperatures appeared to be unrelated to evaporation. Air temperature proved to be insignificant to the evaporation flux, and net radiation alone could only account for 54% of the variability. The combination of the net radiation and the wet and dry bulb temperature depression at 1 m accounted for 88% of the variability of the evaoorative heat flux. The mean α' for a site is assumed to be controlled by the surface type in simplified variations of the combination model. The conclusion has been drawn from this study that the variability of α' can be accounted for by variable atmospheric humidities as well as net radiation. The importance of this atmospheric control on the rate of evaporation is emphasized.</p> / Thesis / Master of Science (MSc)

<p> Plants can control water loss through their stomata in response to changing environmental conditions. Some research has been done on predicting stomatal resistance from climate variables such as solar radiation intensity, temperature, and vapour pressure deficit, but the factors involved are numerous and complex. Information on evapotranspiration from non-cultivated vegetation under field conditions is relatively scarce. This study was carried out in a subarctic coastal wetland on James Bay, an area for which little data exists. Leaf resistance measurements were collected in the field with a diffusion porometer as the 1988 growing season reached its peak. Meteorological data were also recorded. The main species examined was Carex aquatilis, a common sedge with a circumpolar distribution. As has been previously found, field resistances were lower than those reported for greenhouse-grown specimens. Attempts to derive a multivariate regression model to predict leaf resistances had a maximum explained variance under 40%. However, Principal Components Analysis suggested that cool, sunny, low-VPD conditions tend to accompany low leaf resistances. An increase in leaf resistance occurs when solar radiation intensities are low, or when the air is warm and dry. In addition, Carex aquatilis from drier areas showed less growth, and generally higher leaf resistances, than plants growing in saturated soils. Since climate models predict higher temperatures and lower soil moisture for the study area, due to carbon dioxide-induced warming, higher stomatal resistances could result. If these changes are severe enough, primary production could be reduced in the subarctic wetlands. Through the food chain, this could possibly affect species important to the economy and ecology of northern areas.</p> / Thesis / Candidate in Philosophy

Continuous measurements of the energy and radiation balance were made during the 1991 growing season over a dwarf willow-birch forest located near Churchill, Manitoba. The ecological setting is described in terms of both the nature of the substrate and the morphology and distribution of the plant species. Intensive measurements of stomatal conductance and xylem pressure potential for several species were taken on three fair weather days. These represented a wide range of air temperatures and leaf-to-air vapour pressure deficits and allowed the quantification of the surface-atmosphere interactions. The very dynamic and important role of the vegetation in the evaporative process is illustrated. The willow-birch forest consists of six main species which have colonized the recently emerged coastline. There is a wide range in the plant height, rooting networks, and above-to-below ground plant mass. A mature leaf area index of 0.81 m^2 m^-2 was reached within 15 days after the onset of growth. The substrate consists of a 20 cm moderately saline organic layer situated on top of sand. Soil moisture was high, with at least some of the roots of all plants residing within the saturated zone throughout the growing season. The influence of the vegetation on both the radiation and energy balance is illustrated by partitioning the growing season into growth, mature and senescence periods. A strong relationship between surface albedo and vegetation growth indicates that the canopy is more effective in reflecting than in trapping radiation. As the canopy matures, the addition of transpiration to the overall evaporation dramatically increases the magnitude of the latent heat flux at the expense of the sensible heat flux. A sensitivity analysis indicates that evaporation is highly sensitive to the canopy resistance. The sensitivity of evaporation to canopy resistance, in turn, is a function of the ratio of canopy-to-aerodynamic resistance. Strong seasonal and diurnal trends are shown in the sensitivity of evaporation to net radiation, canopy resistance, and aerodynamic resistance. Diurnal stomatal conductance measurements indicate that some species show a pronounced midday stomatal closure. A conceptual model is developed which attributes this behaviour to differences in the sensitivities to the leaf-to-air vapour pressure deficit. A non-linear boundary line analysis of stomatal conductance indicates species-specific responses to irradiance, air temperature, leaf-to-air vapour pressure deficit, and xylem pressure potential. The results of the boundary line analysis are coupled with a modified version of the Penman-Monteith combination model. The model predicts evaporation accurately when the canopy is mature, and indicates that 80% of the evaporation originates from the plants (transpiration). The model is used to examine the potential effects of species composition and climate change on evaporation. This illustrates the important and variable role that vegetation can play in determining responses to climate change. / Thesis / Master of Science (MSc)

This thesis presents results from several studies that have focused on the carbon and nutrient dynamics in soils at the forest – tundra ecotone in Fennoscandia. The main objectives of the study were: (i) to investigate the links between the physical environment, above-ground vegetation communities, soil carbon storage, nutrient status and the chemical composition of the soil organic matter (SOM), and (ii) to quantify trace gas fluxes (methane and carbon dioxide) between mesic soils and the atmosphere. Four main field areas spanning an 8 degree latitudinal gradient were established at the ecotone in 1998 and studied for four years. In addition to the natural gradients we also established a warming treatment. Decomposition rates (i.e. carbon dioxide efflux and litter decomposition) were higher at our forest sites. This was linked principally to the more favourable physical environment at the forest sites, rather than to SOM quality, despite some indications of higher SOM quality at forest sites based upon conventional chemical analysis and 13C NMR techniques. Tundra soils stored large amounts of potentially labile carbon that could readily be accessed by microorganisms when transferred to a forest environment. The interrelation between increased soil temperature and reduced soil moisture content is likely to moderate the response of decomposition rates to increased temperatures. Generally, these mesic soils showed net methane uptake from the atmosphere, which was enhanced by the warming treatment. No differences between forest or tundra soils could be detected. The major conclusions presented here are that (1) soil carbon storage is likely to be reduced if mountain birch forest replaces tundra heath and (2), methane uptake in mesic soils in the Fennoscandian mountains represents a negative feedback to further environmental change in a warmer climate.

Northern lakes and ponds are abundant and emit large amounts of the potent climate forcer methane to the atmosphere at rates prone to change with amplified Arctic warming. In spite of being important, fluxes from surface waters are not well understood. Long-term measurements are lacking and the dominant and irregular transport mode ebullition (bubbling) is rarely quantified, which complicate the inclusion of lakes and ponds in the global methane budget. This thesis focuses on variations in emissions on both local and regional scales. A synthesis of methane fluxes from almost all studied sites constrains uncertainties and demonstrates that northern lakes and ponds are a dominant source at high latitudes. Per unit area variations in flux magnitudes among different types of water bodies are mainly linked to water depth and type of sediment. When extrapolated, total area is key and thus post-glacial lakes dominate emissions over water bodies formed by peat degradation or thermokarst processes. Further, consistent multiyear measurements in three post-glacial lakes in Stordalen, northern Sweden, reveal that seasonal ebullition, primarily driven by fermentation of acetate, can be predicted by easily measured parameters such as temperature and heat energy input over the ice-free season. Assuming that most water bodies respond similarly to warming, this thesis also suggests that northern lakes and ponds will release substantially more methane before the end of the century, primarily as a result of longer ice-free seasons. Improved uncertainty reductions of both current and future estimates rely on increased knowledge of landscape-level processes related to changes in aquatic systems and organic loading with permafrost thaw, as well as more high-quality measurements, seldom seen in contemporary data. Sampling distributed over entire ice-free seasons and across different depth zones is crucial for accurately quantifying methane emissions from northern lakes and ponds. / <p>At the time of the doctoral defense, the following paper was unpublished and had a status as follows: Paper 4: Manuscript.</p>

Climate change is expected to continue to cause large increases in temperature in Arctic and sub-Arctic ecosystems which has already resulted in changes to plant communities; for example, increased shrub biomass and range. It is important to understand how warmer temperatures could affect the plant community in a wetland system because this region provides crucial high-quality forage for migratory herbivores during the breeding season. One mechanism by which warming could cause change is directly, where warming influences the vital rates of a species; these effects may be either positive or negative. Warmer temperatures may also affect a species indirectly, by impacting neighboring plants which compete with, or facilitate that species. Altering interspecific interactions may affect the abundances of the surrounding species. Recent research shows these ‘indirect’ effects which are mediated by biotic interactions may be important enough to reverse ‘direct’ effects of climate change in some plant communities. Furthermore, herbivores have been shown to mediate the effects of warming, in some systems, even preventing shrub expansion. However, the abundance of herbivores may change because of climate change so it is important to understand the role of herbivores in mitigating climate change effects to inform management strategy. Therefore, we aimed to determine the importance of direct and indirect effects of warming on this plant community while considering changing herbivore pressures. We conducted a two-year field experiment in the coastal wetlands of western Alaska to investigate how warming and herbivory will impact the abundances of two common species, a sedge and a dwarf shrub. We used the results from the experiment to predict the equilibrium abundances of the two species under different climate and herbivory scenarios and determine the contribution of direct and indirect effects to predicted community change. The sedge, Carex ramenskii, remained dominant in under ambient conditions, but the dwarf shrub, Salix ovalifolia, became dominant in warmed treatments. Herbivory mediated some of the effects of warming; where grazing was present community composition did not change as much as where it was not grazed. Results suggest that in the absence of goose herbivory, a 2°C increase could cause a shift from sedge to woody plant dominance on the coast of western Alaska. However, if grazing pressure by geese continues at the present rate, it may help retain the current community composition, though herbivory pressure was not sufficient to entirely reverse the effect of warming. Finally, we found that direct effects were more important than indirect effects in causing changes to this plant community.

<p>This thesis presents results from several studies that have focused on the carbon and nutrient dynamics in soils at the forest – tundra ecotone in Fennoscandia. The main objectives of the study were: (i) to investigate the links between the physical environment, above-ground vegetation communities, soil carbon storage, nutrient status and the chemical composition of the soil organic matter (SOM), and (ii) to quantify trace gas fluxes (methane and carbon dioxide) between mesic soils and the atmosphere. Four main field areas spanning an 8 degree latitudinal gradient were established at the ecotone in 1998 and studied for four years. In addition to the natural gradients we also established a warming treatment. Decomposition rates (i.e. carbon dioxide efflux and litter decomposition) were higher at our forest sites. This was linked principally to the more favourable physical environment at the forest sites, rather than to SOM quality, despite some indications of higher SOM quality at forest sites based upon conventional chemical analysis and ¹³C NMR techniques. Tundra soils stored large amounts of potentially labile carbon that could readily be accessed by microorganisms when transferred to a forest environment. The interrelation between increased soil temperature and reduced soil moisture content is likely to moderate the response of decomposition rates to increased temperatures. Generally, these mesic soils showed net methane uptake from the atmosphere, which was enhanced by the warming treatment. No differences between forest or tundra soils could be detected.</p><p>The major conclusions presented here are that (1) soil carbon storage is likely to be reduced if mountain birch forest replaces tundra heath and (2), methane uptake in mesic soils in the Fennoscandian mountains represents a negative feedback to further environmental change in a warmer climate.</p>