Since the late 1880's western juniper has expanded in range and increased in density in sagebrush-bunchgrass, riparian, and forested plant communities of the Pacific Northwest. Succession to western juniper woodland has been shown to reduce the productivity and diversity of the understory component, result in concentration of soil nutrients beneath juniper canopies, and reduces soil moisture storage. This study assessed understory plant succession, soil nitrogen (N) cycling, litter decomposition, and soil moisture availability following cutting of a western juniper woodland on Steens Mountain, Oregon. Cutting of western juniper reduced below-ground competition for water and N, increasing soil moisture storage and N availability for understory species. Leaf water potentials were less negative, and N concentration and biomass in understory plants were greater in the cut treatment. Understory species responded to improved growth conditions with increased cover, biomass, density, and diversity. In 1993, total understory biomass and canopy cover were 870% and 300% greater, respectively, in the cut treatment than the uncut woodland. Understory succession was dominated by plants present on the site prior to cutting. Results indicated initial that bunchgrass densities of 2 plants/m2 were sufficient for perennial grasses to dominate following juniper cutting. Juniper cutting is an effective method for restoring the understory component in sagebrush rangelands that are currently dominated by western juniper woodland. Nitrogen availability was greatest in cut-interspace locations the first year following treatment and in cut-duff locations in the second year. Nitrification was lowest in cut-slash and woodland-duff locations, areas receiving fresh litter inputs and experiencing lower temperatures than interspace (cut and woodland treatments) and cut-duff locations. Decomposition of juniper litter was two times faster in the cut treatment, however the release of litter N occurred earlier in the woodland. Large inputs of N poor litter from cut juniper slash were hypothesized to have increased microbial demand for N, resulting in immobilization of litter N. Immobilization of litter N may be important in conserving N on sites following cutting. / Graduation date: 1997

Graduation date: 1995

Forests are the dominant terrestrial ecosystem in Ontario and were once widespread throughout much of eastern North America; in southern Ontario, forest cover has declined from 90% to approximately 17% in the past 200 years. Conversion of the surrounding landscape to urban and agricultural land uses creates edge effects which increase the impact of forest cover loss and threaten ecological integrity. Other impacts on forest ecological integrity include incompatible human activities, a generally inhospitable landscape matrix, and microenvironment changes, including those from chemical pollution and heat island effects. Coupled with direct degradation from human activities like free-riding, the altered forest microenvironment can facilitate increased invasion by opportunistic and competitive plant species, e.g. Alliaria petiolata (Brassicaceae). This tends to be worse in urban areas because of greater visitation that creates soil compaction, trampling, and more vectors to introduce the seeds of invasive species. Active management is often necessary in areas which have been degraded. Ecological restoration represents an important active management tool for urban forest protected area managers. The ecological restoration of the forest understory in altered and threatened habitats represents an important aspect of forest restoration. For this study, I tested the effectiveness of an ecological restoration using a multi-species assemblage approach with three native understory species to restore the herbaceous layer in Natchez Hills, an Environmentally Sensitive Policy Area in Kitchener, Ontario. The ecological restoration site was degraded and this reduced native plant species in the forest understory. The species selected, Caulophyllum giganteum (Farwell) Loconte & W.H. Blackwell (Berberidaceae), Podophyllum peltatum Linnaeus (Berberidaceae) and Maianthemum racemosum subsp. racemosum (Linneaus) Link (Asparagaceae), are widespread throughout eastern North America, and were selected for use in this study because of their persistence and relative abundance in Natchez Hills ESPA. Adult transplants were planted into 2 metre by 2 metre quadrats in different densities and shoot populations were monitored for two field seasons to determine if density played a role in early ecological restoration success. The methodology I designed for the ecological restoration was as practical as possible, given the real world financial and logistical constraints faced by many urban forest managers. Statistical analysis using repeated measures multivariate analysis of variance showed aerial shoot density did not have a significant effect on early restoration success rates (p > 0.05). However, there was a significant increase in restored species. Caulophyllum giganteum and Podophyllum peltatum shoot populations increased by over 97% at the ecological restoration site one year after transplanting. Maianthemum racemosum subsp. racemosum shoot populations increased by 51% at the site. For comparison, another study in the same forest reported early ecological restoration rates of 92% to 100% for herbaceous understory species, including C. giganteum and P. peltatum. Other North American studies using transplanted perennial herbaceous material reported average survival rates of 16.6% two years after planting (varying from 1.25% to 51.4% using eight herbaceous species in the Boston area), to 91% one year after restoration (varying from 42.1% to 100% using 19 herbaceous species of the deciduous forest in the Midwest). In instances where protected areas are degraded, active management and ecological restoration are often necessary tools. This ecological restoration demonstrated that the native herbaceous species used in this study are suitable for use in early ecological restorations in maple beech upland forests in the City of Kitchener. Using a practical and cost-effective methodology this ecological restoration experiment showed that it is possible to implement urban restoration of the forest understory in degraded areas, and to achieve success in early ecological restoration using native herbaceous forest understory species.

Thousands of hectares of red pine (Pinus resinosa Ait.) plantations were established during the early 20th century to restore abandoned agricultural lands and increase forest cover. Concern over the ecological integrity [i.e. the degree to which a community is determined to be characteristic of its natural region, including abiotic and biotic components, likely to persist and maintain the rates of change and supporting processes (Canada National Parks Act (S.C. 2000, c. 32) 2011) of these plantations and a desire to increase native forest cover has prompted a more thorough understanding of the understory succession of these plantations. This study addresses the question, how does the understory vegetation composition and diversity in managed red pine plantations compare to the understory of a reference deciduous forest?. I examined the understory diversity in a chronosequence of managed red pine plantations owned by Simcoe County. Biophysical characteristics including diffuse non-interceptance (DIFN), tree regeneration, litter depth, and soil physical and chemical properties were gathered from three replicates of each of four age groups and from a reference deciduous forest group in order to determine how each variable changed and influenced the succession of the understory diversity. Field investigations were completed during the spring, summer, and fall of 2009 in order to examine seasonal differences in diversity. A two-way analysis of variance with a Tukey’s post hoc test was used to determine if the forest groups were significantly different from each other for each variable measured. Species diversity (measured by the Shannon-Wiener Index) generally increased with plantation age and seasonal differences were apparent. A lack of significant difference between the oldest plantation group (est. 1927-1932) and the reference forest group indicates diversity is increasing; this is intriguing because, in contrast, species composition is only 30% similar. This may indicate either not enough time has passed for the understory to regenerate or the understory in the plantations is succeeding in a different trajectory. Furthermore, spring ephemeral species were lacking in the understory of the plantation forests indicating this phenological group is not regenerating in the plantations. Tree density is generally increasing in the plantation groups and the dominant tree species regenerating in the oldest plantation group includes a similar suite of species to that found in the reference forest groups. This indicates that the canopy is succeeding towards a native forest community. Pearson correlation analysis revealed that litter depth is significantly negatively correlated to diversity in the summer and fall (p<0.01). Litter depth was also the primary predictor variable produced by the forward stepwise regression in the summer and fall indicating that litter depth is a major driver of diversity. From comparative studies, a deeper litter layer often inhibits the germination and emergence of herbaceous species; from my study, this may explain the lack of spring ephemerals in the plantation groups observed. DIFN was significantly negatively correlated to diversity in the spring (p<0.01) and was the predictor variable for spring diversity indicating that light characteristics in the spring are a major variable influencing spring diversity. With increasing age of plantation group the DIFN followed a similar pattern to the reference forest group indicating that the thinning regime and regeneration is shifting the light characteristics to that found in a natural forest stand. I conclude that the thinning regime in red pine plantation is promoting the regeneration of trees and increasing the diversity of understory vegetation. However, in order to direct the successional trajectory of the understory vegetation of red pine plantations to a composition similar to the native woodlands of that area, a modified management program should be considered and active restoration should be implemented.

No description available.

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

Forests are the dominant terrestrial ecosystem in Ontario and were once widespread throughout much of eastern North America; in southern Ontario, forest cover has declined from 90% to approximately 17% in the past 200 years. Conversion of the surrounding landscape to urban and agricultural land uses creates edge effects which increase the impact of forest cover loss and threaten ecological integrity. Other impacts on forest ecological integrity include incompatible human activities, a generally inhospitable landscape matrix, and microenvironment changes, including those from chemical pollution and heat island effects. Coupled with direct degradation from human activities like free-riding, the altered forest microenvironment can facilitate increased invasion by opportunistic and competitive plant species, e.g. Alliaria petiolata (Brassicaceae). This tends to be worse in urban areas because of greater visitation that creates soil compaction, trampling, and more vectors to introduce the seeds of invasive species. Active management is often necessary in areas which have been degraded. Ecological restoration represents an important active management tool for urban forest protected area managers. The ecological restoration of the forest understory in altered and threatened habitats represents an important aspect of forest restoration. For this study, I tested the effectiveness of an ecological restoration using a multi-species assemblage approach with three native understory species to restore the herbaceous layer in Natchez Hills, an Environmentally Sensitive Policy Area in Kitchener, Ontario. The ecological restoration site was degraded and this reduced native plant species in the forest understory. The species selected, Caulophyllum giganteum (Farwell) Loconte & W.H. Blackwell (Berberidaceae), Podophyllum peltatum Linnaeus (Berberidaceae) and Maianthemum racemosum subsp. racemosum (Linneaus) Link (Asparagaceae), are widespread throughout eastern North America, and were selected for use in this study because of their persistence and relative abundance in Natchez Hills ESPA. Adult transplants were planted into 2 metre by 2 metre quadrats in different densities and shoot populations were monitored for two field seasons to determine if density played a role in early ecological restoration success. The methodology I designed for the ecological restoration was as practical as possible, given the real world financial and logistical constraints faced by many urban forest managers. Statistical analysis using repeated measures multivariate analysis of variance showed aerial shoot density did not have a significant effect on early restoration success rates (p > 0.05). However, there was a significant increase in restored species. Caulophyllum giganteum and Podophyllum peltatum shoot populations increased by over 97% at the ecological restoration site one year after transplanting. Maianthemum racemosum subsp. racemosum shoot populations increased by 51% at the site. For comparison, another study in the same forest reported early ecological restoration rates of 92% to 100% for herbaceous understory species, including C. giganteum and P. peltatum. Other North American studies using transplanted perennial herbaceous material reported average survival rates of 16.6% two years after planting (varying from 1.25% to 51.4% using eight herbaceous species in the Boston area), to 91% one year after restoration (varying from 42.1% to 100% using 19 herbaceous species of the deciduous forest in the Midwest). In instances where protected areas are degraded, active management and ecological restoration are often necessary tools. This ecological restoration demonstrated that the native herbaceous species used in this study are suitable for use in early ecological restorations in maple beech upland forests in the City of Kitchener. Using a practical and cost-effective methodology this ecological restoration experiment showed that it is possible to implement urban restoration of the forest understory in degraded areas, and to achieve success in early ecological restoration using native herbaceous forest understory species.

Thousands of hectares of red pine (Pinus resinosa Ait.) plantations were established during the early 20th century to restore abandoned agricultural lands and increase forest cover. Concern over the ecological integrity [i.e. the degree to which a community is determined to be characteristic of its natural region, including abiotic and biotic components, likely to persist and maintain the rates of change and supporting processes (Canada National Parks Act (S.C. 2000, c. 32) 2011) of these plantations and a desire to increase native forest cover has prompted a more thorough understanding of the understory succession of these plantations. This study addresses the question, how does the understory vegetation composition and diversity in managed red pine plantations compare to the understory of a reference deciduous forest?. I examined the understory diversity in a chronosequence of managed red pine plantations owned by Simcoe County. Biophysical characteristics including diffuse non-interceptance (DIFN), tree regeneration, litter depth, and soil physical and chemical properties were gathered from three replicates of each of four age groups and from a reference deciduous forest group in order to determine how each variable changed and influenced the succession of the understory diversity. Field investigations were completed during the spring, summer, and fall of 2009 in order to examine seasonal differences in diversity. A two-way analysis of variance with a Tukey’s post hoc test was used to determine if the forest groups were significantly different from each other for each variable measured. Species diversity (measured by the Shannon-Wiener Index) generally increased with plantation age and seasonal differences were apparent. A lack of significant difference between the oldest plantation group (est. 1927-1932) and the reference forest group indicates diversity is increasing; this is intriguing because, in contrast, species composition is only 30% similar. This may indicate either not enough time has passed for the understory to regenerate or the understory in the plantations is succeeding in a different trajectory. Furthermore, spring ephemeral species were lacking in the understory of the plantation forests indicating this phenological group is not regenerating in the plantations. Tree density is generally increasing in the plantation groups and the dominant tree species regenerating in the oldest plantation group includes a similar suite of species to that found in the reference forest groups. This indicates that the canopy is succeeding towards a native forest community. Pearson correlation analysis revealed that litter depth is significantly negatively correlated to diversity in the summer and fall (p<0.01). Litter depth was also the primary predictor variable produced by the forward stepwise regression in the summer and fall indicating that litter depth is a major driver of diversity. From comparative studies, a deeper litter layer often inhibits the germination and emergence of herbaceous species; from my study, this may explain the lack of spring ephemerals in the plantation groups observed. DIFN was significantly negatively correlated to diversity in the spring (p<0.01) and was the predictor variable for spring diversity indicating that light characteristics in the spring are a major variable influencing spring diversity. With increasing age of plantation group the DIFN followed a similar pattern to the reference forest group indicating that the thinning regime and regeneration is shifting the light characteristics to that found in a natural forest stand. I conclude that the thinning regime in red pine plantation is promoting the regeneration of trees and increasing the diversity of understory vegetation. However, in order to direct the successional trajectory of the understory vegetation of red pine plantations to a composition similar to the native woodlands of that area, a modified management program should be considered and active restoration should be implemented.

Boreal mixedwood forest stands are comprised of a mixture of small canopy patches of varying dominance by conifer (mostly white spruce (Picea glauca (Moench) Voss)) and broadleaf (mostly trembling aspen (Populus tremuloides Michx.) trees. The purpose of this work was to extend our understanding of the patterns and causes of variation in understory vascular plant communities in unmanaged, mature boreal mixedwood forests. First, I assessed variation in understory community composition in relation to canopy patch type (conifer, mixed conifer-broadleaf, broadleaf, gaps) within mixedwood stands. The mosaic of canopy patches leads to different micro-habitat conditions for understory species, allowing for communities that include both early and late successional species and contributing to greater understory diversity. This study suggests that the mosaic of small canopy patches within mixed forest stands resembles a microcosm of the boreal mixedwood landscape, across which understory community composition varies with canopy composition at the stand scale. Second, I investigated the hierarchical organization of understory diversity in relation to the heterogeneous mosaic of canopy patch types through additive partitioning of diversity. The largest proportion of species richness was due to turnover among patches within patch type while individual patches had higher evenness. The mosaic of canopy patch types within mixedwood forests likely plays a crucial role in maintaining the hierarchical levels at which understory diversity is maximized. Third, I examined interactions among understory plant species by investigating the effect of shrub removal on biomass, composition and diversity of herbs using a 3-yr removal study in a natural understory community. There is asymmetric competition for light between erect shrub and herb species but herb response to erect shrub removal was species-specific. Plant interactions play an important role in structuring boreal understory communities. Finally, I explored the relative influence of space, environmental variables, and their joint effects, on understory composition and richness. The environmental variation caused by small canopy patches and biotic processes, such as species interactions, converge at the fine scale to create a spatially patchy structure in understory communities in boreal mixedwood forests. Modifications in the natural mixture of small canopy patches could disrupt the spatial and environmental structures that shape understory composition and diversity patterns. / Forest Biology & Management

Boreal mixedwood forest stands are comprised of a mixture of small canopy patches of varying dominance by conifer (mostly white spruce (Picea glauca (Moench) Voss)) and broadleaf (mostly trembling aspen (Populus tremuloides Michx.) trees. The purpose of this work was to extend our understanding of the patterns and causes of variation in understory vascular plant communities in unmanaged, mature boreal mixedwood forests. First, I assessed variation in understory community composition in relation to canopy patch type (conifer, mixed conifer-broadleaf, broadleaf, gaps) within mixedwood stands. The mosaic of canopy patches leads to different micro-habitat conditions for understory species, allowing for communities that include both early and late successional species and contributing to greater understory diversity. This study suggests that the mosaic of small canopy patches within mixed forest stands resembles a microcosm of the boreal mixedwood landscape, across which understory community composition varies with canopy composition at the stand scale. Second, I investigated the hierarchical organization of understory diversity in relation to the heterogeneous mosaic of canopy patch types through additive partitioning of diversity. The largest proportion of species richness was due to turnover among patches within patch type while individual patches had higher evenness. The mosaic of canopy patch types within mixedwood forests likely plays a crucial role in maintaining the hierarchical levels at which understory diversity is maximized. Third, I examined interactions among understory plant species by investigating the effect of shrub removal on biomass, composition and diversity of herbs using a 3-yr removal study in a natural understory community. There is asymmetric competition for light between erect shrub and herb species but herb response to erect shrub removal was species-specific. Plant interactions play an important role in structuring boreal understory communities. Finally, I explored the relative influence of space, environmental variables, and their joint effects, on understory composition and richness. The environmental variation caused by small canopy patches and biotic processes, such as species interactions, converge at the fine scale to create a spatially patchy structure in understory communities in boreal mixedwood forests. Modifications in the natural mixture of small canopy patches could disrupt the spatial and environmental structures that shape understory composition and diversity patterns. / Forest Biology & Management