This study examines the influence of climate on tree-ring properties of several major tree species: Pacific silver fir (Abies amabilis (Dougl. ex Loud.) Forbes), subalpine fir (Abies lasiocarpa (Hook.) Nutt.), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), and lodgepole pine (Pinus contorta var. latifolia Dougl. ex Loud.). Our three objectives were to determine how (1) tree-ring properties change along an elevation gradient, (2) short-term climatic influences are correlated with tree-ring properties, and (3) long-term climatic influence on tree-ring properties.

The Mountain Hemlock (MH) zone includes all subalpine forests along British Columbia’s coast. It occurs at elevations where most precipitation falls as snow and the growing season is less than 4 months long. The zone includes the continuous forest of the forested subzones and the tree islands of the parkland subzones (Figure 1). Old-growth stands are populated by mountain hemlock, Pacific silver fir, and Alaska yellow-cedar, and are among the least-disturbed ecosystems in the world. Canopy trees grow slowly and are commonly older than 600 years, while some Alaska yellow-cedars may be up to 2000 years old. Understanding regeneration patterns in the MH zone has become increasingly important as logging continues towards higher elevations of the zone where snowpacks are deeper.

Height growth models of coastal low- and mid-elevation Pacific silver fir, low-elevation white spruce, and low- and midelevation lodgepole pine have been used for predicting productivity of subalpine fir, Engelmann spruce, and lodgepole pine, respectively. These models, however, are biased in predicting height growth of high-elevation subalpine fir, Engelmann spruce, and lodgepole pine. To improve this situation, 329 sample plots (165 for subalpine fir, 90 for Engelmann spruce, and 74 for lodgepole pine) were located throughout the Engelmann Spruce-Subalpine Fir (ESSF) zone. Stem analysis was carried out on three dominant trees in each 0.04 ha sample plot. For each study species, a height growth model was developed on the data from two-thirds of the sample plots using the conditioned Chapman-Richards’ function; the model was validated using the remaining one-third of the sample plots.

Subalpine ecosystems are experiencing rapid changes in snow pack, temperature, and precipitation regime as a result of anthropogenic climate forcing. These changes in climate can have a profound effect on subalpine ecosystem structure and functioning, which may ultimately feed back to climate change. In this study, I examined the response of the subalpine meadow plant communities at the Rocky Mountain Biological Laboratory to natural and simulated climate change. First, I looked at whether changes in growing season precipitation or temperature regime would have the larger effect on subalpine ecosystem carbon flux. In a simulated warming experiment, changes in growing season precipitation had a tenfold larger effect on cumulative carbon flux than did the warming treatment. Along a natural climatic and elevational gradient, precipitation stimulates carbon uptake, particularly at higher elevations. Given projected decreases in summer precipitation in the high elevation Rockies, we predict a 20% decrease in carbon uptake from subalpine meadows. Second, I compared the taxonomic, phylogenetic and functional structure of plant communities along an elevational gradient to infer which climatic and biotic factors influence community assembly at each elevation. Floral and phenology traits become overdispersed at high elevation, mirroring phylogenetic relatedness, and suggesting pressure to diversify to attract pollinators during the abbreviated growing season. At the same time, leaf functional traits become clustered at high elevation, indicating multiple opposing assembly mechanisms in subalpine communities. Finally, I studied the natural history of sagebrush, Artemisia tridentate ssp. vaseyana, at its elevational range limit in subalpine meadows. In particular, I focused on the importance of warming and species interactions in elevational advance of the species. I found that facilitation by neighboring forbs was critical for sagebrush seedling survival, decreasing mortality by 75%. Seedling mortality was overwhelmingly due to desiccation of seedlings; therefore, neighboring forbs moderate temperature and water stress for seedlings. Despite the extremely limited growing season at high elevation caused by subfreezing temperatures, subalpine ecosystem structure and function are closely tied to water availability during the growing season. Therefore, improved predictions of future precipitation regimes over the Rocky Mountains will be our best tool for conservation of these fragile habitats.

Le massif de Sixt constitue la couverture sedimentaire de la retombée Ouest du massif cristallin des Aiguilles Rouges . A ce titre, il occupe dans la zone externe des Alpes, la même place que le Vercors, la Chartreuse , les Bauges ou les Aravis. Pourtant, en se déplaçant du Sud au nord dans la zone subalpine, on constate une complication des structures et une tendance à la tectonique souple . Insensiblement la couverture décollée et plissée des environs de Grenoble va passer à des structures d 'amplitude plus forte pour aboutir aux nappes helvétiques . Par sa position, le massif de Sixt semble , alors , devoir prendre un caractère propre : celui de zone de transition . Les travaux de J.ROSSET sur la chaîne des Aravis (1957) ont montré que la tectonique de cette région s 'expliquait t r è s bien sans faire intervenir la notion de nappe : or , et nous l e verrons par la suite , j 'ai pu faire de nombreuses constatations analogues à celles de J .ROSSET . Pour l ' ensemble du massif de Sixt , plusieurs problèmes se sont posés : fallait - il toujours considérer les plis du Jurassique supérieur du Fond de la Combe comme eprésentant le flanc inverse d ' une nappe, et la "fenêtre du Pelly" comme un témoin de l'autochtone? Mon collègue X.PIERRE travaillant plus au Sud, s'est heurté aux mêmes difficultés: son étude s'est orientée sur l a "fenêtre des Fonts" et sur la présence d 'un éventuel Jurassique au contact du Cristallin des Aiguilles Rouges près des chalets de Moëde,

Quantifying carbon emissions of water bodies at regional scale is required as recent studies revealed their contribution in carbon cycling is significant. This demands to scale up water bodies carbon emissions from local to regional scale using as accurate approach as possible. In this study data of carbon (CO2-C) fluxes for 80 sampled lakes were used to scale up to more than 3000 lakes and ponds over the catchment. The most appropriate method for upscaling was the one in which two factors of water body size and location (altitude) were involved and the uncertainties were quantified in an advanced approach (Monte Carlo model). Based on the estimates obtained in this method, the annual carbon emission from all water bodies (~ 500 km2) was about 2900 ton C yr-1 . About 62% of this annual emission was related to the large lake Torneträsk (334 km2) and another 38% to all other lakes and ponds (166 km2). Water bodies in subalpine region dominated (90%) total water bodies area and were the major contributor (97%) to the total carbon emissions of all water bodies. The remaining small contribution (3%) was for water bodies in the alpine region, which contains only 10% of total water bodies area. These data indicate that all water bodies smaller than the large lake Torneträsk especially the ones in the subalpine region have considerable contribution to the annual carbon emission of all water bodies. Considering water body size and altitude factors in the advanced upscaling method was appropriate to obtain accurate estimates.

Whitebark pine (Pinus albicaulis) is an endangered keystone tree species growing at the highest elevations in the mountain ranges of western North America. Across its range, whitebark pine is faced with a number of threats including fire suppression, mountain pine beetle, white pine blister rust, and climate change. Climate change is perhaps the greatest threat facing the species, yet it is the least understood. Most studies rely on model predictions and only look at the impacts on whitebark pine itself, not taking into consideration the other bird, mammal, and plant communities that are associated with it. In order to assess the potential effects of climate change on whitebark pine communities in the southern Coast Mountains of British Columbia, this thesis examined the vegetation associations and climate controls currently shaping the communities. My results showed that whitebark pine is growing in the open away from other subalpine tree species. This suggests that whitebark pine is not facilitating other subalpine tree species, contrary to what has been shown in the Rocky Mountains. Evidence of a distinct suite of understory vegetation associated with whitebark pine is weak and inconclusive. Differences in understory vegetation appear to be mainly due to site differences in climate, soils, and topography. Age distributions constructed from tree cores revealed that whitebark pine decline at lower elevation sites may be due to successional advancement to subalpine fir, and subalpine fir is currently encroaching into higher elevation sites. A dendrochronological assessment revealed that winter conditions, including snowpack, temperature, and the Aleutian Low Pressure Index (ALPI) were the most limiting to whitebark pine growth at high-elevation sites, but biotic factors including disease and competition appear to be more important than climate in determining annual ring growth at lower elevation sites. Bootstrapped correlations between annual ring widths and snowpack records showed that tree responses to fluctuating snowpack have changed over time. For most of the 20th century, low snowpack periods were associated with greater annual growth. Since around 1970, when the snowpack levels dropped below anything previously recorded for the area, annual tree growth has been reduced. It appears that these high elevation tree species require a balance between too much snow (shorter growing season) and too little snow (reduced protection from harsh winter conditions). Climate change models for the area predict drastically reduced snowpack in the coming decades. If snowpack continues to drop, as it has since 1970, it will likely lead to severe impacts on whitebark pine growth in the southern Coast Mountains. / Graduate / 0329 / carlsonkim@hotmail.com

The Bogong High Plains are a series of alpine and subalpine plateaux in NE Victoria. The vegetation of the High Plains consists of woodland, heathland, grassland herbfield and wetland communities. On the better drained sites, the transition from heath dominated communities to grass dominated communities is correlated with gradients of decreasing accumulation and persistence of snow, increasing exposure to wind and low minimum air temperatures, and decreasing steepness and rockiness of terrain. In many areas, shrubs have increased in cover and abundance, especially over the last 50 years. In particular, shrubs have invaded many areas of grassland, which has resulted in an expansion of both open heath and closed heath. The High Plains have been grazed by free ranging cattle each summer since the 1850's, and a primary aim of this thesis is to investigate the dynamics of heathland and grassland, and the impact that cattle grazing has upon these two vegetation types.

Subalpine larch (Larix lyallii Parl.) has a narrow ecological niche at timberline in the Cascade Range and the Rocky Mountains of western North America. Demographic factors, including a long generation time (average 500 years) and a late arrival at sexual maturity (100-200 years), make it unlikely that this species will be able to adapt to predicted climate change. A better understanding of genetic structure and genetic diversity is necessary in order to effectively manage this species for future generations. Foliage from 62 populations of subalpine larch was collected in order to elucidate the range-wide population genomics of the species. DNA was extracted and a next-generation sequencing method, restriction site associated DNA sequencing (RAD-seq), was used to generate genome-wide single nucleotide polymorphism (SNP) marker data. Three genetically differentiated clusters were identified via principal components analysis, a discriminant analysis of principal components and Bayesian STRUCTURE analysis: the Cascade Range, the southern Rocky Mountains and the northern Rocky Mountains. A monophyletic group in the central Rocky Mountains was also identified in a dendrogram of genetic distance but this group had weak bootstrap support (49%), meaning genetic differentiation depends on relatively few genetic variants. Genetically differentiated groups should be prioritized for future management and conservation efforts. Negative values of Tajima’s D and preferred demographic scenarios generated by coalescent simulations indicated that 15 populations all have a recent history of expansion. Genetic diversity within these populations was found to be moderate (HO = 0.15 – 0.20), inbreeding coefficients were found to be high (FIS = 0.15 – 0.25) and genetic differentiation among populations was found to be high (average FST = 0.18). These results indicated that fragmentation driven by Holocene warming may have resulted in reduced effective population sizes. Smaller populations experience stronger genetic drift and an increased likelihood of inbreeding, which may hinder an adaptive response to natural selection. Still, parameter estimates for preferred demographic scenarios suggested a minimum effective population size of around 20,000 individuals, which is not considered small by most conservationists. A final study of 18 populations found local adaptation to cold temperature in the northern portion of the species range. In all seasons, populations from the northern Rocky Mountains had significantly higher cold tolerance than populations from the central Canadian Rocky Mountains and the northern Cascades. Winter cold tolerance showed strong clines associated with the frost-free period and degree days below zero. These two climate variables explained 65% of the explainable variance in phenotype when redundancy analysis models were conditioned on geography. Seven SNPs were identified that explained a significant portion of the variance in winter cold tolerance. Range-wide, additional SNPs were identified as FST outliers and/or as significantly correlated with environmental gradients, even after correcting for neutral genetic structure. Together, the results of this work indicate that dispersal, neutral evolutionary processes and natural selection have all played important roles in shaping patterns of genetic variation across the natural range of subalpine larch. All of these factors should be considered during the development of management and conservation strategies for this high-elevation conifer species. / Graduate

Les dépôts azotés (DA) mesurés à l'étage subalpin des Pyrénées sont compris entre 8 et 15 kg N ha-1 an-1, soit égaux ou supérieurs à la charge critique actuellement définie pour ces milieux et supérieurs aux estimations fournies par les modèles de chimie-transport. Au cours des soixante dernières années, les pelouses subalpines pyrénéennes sont devenues plus thermophiles et mésotrophes, plus riches en espèces et plus homogènes à l'échelle des Pyrénées. Le changement climatique (CC) et le cumul de DA ont réduit les contraintes abiotiques de ces habitats, favorisant l'installation d'espèces moins stress-tolérantes. Ces milieux piègent et conservent efficacement l'azote d'origine atmosphérique, même en contexte de CC. Des effets des DA sont détectables sur la végétation (%N, biomasse et recouvrement de certains groupes) avant d'être détectables dans le sol. Ces effets sont plus marqués en réponse aux apports dominés par l'ammonium plutôt que par le nitrate, et sont additifs ou compensatoires avec ceux du CC, mais rarement interactifs. / Atmospheric nitrogen deposition in the subalpine Pyrenees is in order of 8 to 15 kg N ha-1 yr-1, equivalent or higher than the actual critical load for these habitats and higher than chemistry-transport models' estimates. During the past six decades, Pyrenean subalpine grasslands became more thermophilous and mesotrophic, richer in species and more homogeneous at the regional scale. Climate change and cumulative nitrogen deposition reduced the abiotic constraints in these habitats, favouring the installation of less stress-tolerant species. These habitats trap and store efficiently nitrogen from deposition, even in a climate change context. Effects are visible in the vegetation (% of nitrogen, biomass and cover of some groups) before being visible in the soil. These effects are more pronounced following ammonium than nitrate dominated additions, and are additive or compensatory with the effects of climate change, but rarely interactive.