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There has been an increased interest in characterizing and interpreting ecological heterogeneity over space and time in the past two decades. This is mainly due to the renewed recognition of the significance of heterogeneity in ecological theories. However, studies that have combined both spatial and temporal aspects of heterogeneity have been rare. A unified approach to define and quantify heterogeneity has also been lacking. Designed to overcome these problems, this study was conducted in a 40-year-old Pinus elliottii Engelm. forest at the Savannah River Site near Aiken, SC, USA with the following specific objectives: 1) to characterize the spatial patterns of soil and forest floor variables (moisture, pH, soil available nitrogen and phosphate, forest floor and soil carbon and nitrogen), 2) to examine the dynamics of these spatial patterns in response to two types of disturbance: whole-tree harvesting and girdling, and 3) to evaluate some of the current methods for quantifying ecological heterogeneity. In response to both disturbance treatments, spatial heterogeneity measured by sample variance showed a marked "increase and then decline" temporal pattern in soil moisture, soil available nitrogen and phosphorus. Similar patterns were not found in total soil C and N, and total litter C and N. Harvesting resulted in greater and more drastic changes in the variations of soil nutrients and water than did girdling. Despite the popularity of semivariogram analysis in recent ecological studies, the technique did not provide consistent results on patterns of heterogeneity in our system. A simulation experiment demonstrated that semivariogram analysis may suffer from many problems when it is used to characterize patchiness, one form of heterogeneity. The results from this study have a number of implications. First, spatial patterns of soil resources are high dynamic. The dynamics of patterns in soil resources may partly account for the weak correlation between vegetation and soil observed in ecological literature. Second, heterogeneity may be most effectively quantified by first identifying quantifiable components and then quantifying these components individually. A common pattern can be sought by comparing patterns of different components of heterogeneity for a given ecological property and by comparing patterns of different ecological variables for a given component of heterogeneity. Third, compared to surveys, field manipulative experiments can provide information that link patterns with ecological processes. As such, this study adds to ecological literature valuable information on temporal changes of soil heterogeneity following disturbance and conceptual advances in the quantification of ecological heterogeneity. Ph. D.

Chapter 1 Abstract: We examined the impact of sampling interval on fine root production and mortality estimates by comparing data from a weekly minirhizotron sampling regimen to subsets of the same data representing biweekly, monthly, bimonthly, and quarterly sampling regimens. We also investigated possible sources of error involved in the root tracing technique and estimated root herbivory using the full weekly sampling regimen. Data were collected for eleven months from a Pinus palustris Miller woodland in southwest Georgia. As sampling interval increased, estimates of production and mortality declined, while estimates of mean fine root lifespan increased. Annual production values ranged from a maximum of 1.26 mm/cm2 for weekly sampling to 0.83 mm/cm2 for quarterly sampling. Total mortality varied from 0.97 mm/cm2 to 0.53 mm/cm2. Bias increased at a decreasing rate when sample interval was increased from weekly to monthly. The root tracing protocol added some small, random error to growth measurements; re-measuring roots returned values 0.16% smaller than initial measures. We also observed a root mortality and regrowth phenomenon that may be measurement error or short-term fluctuation in root length. Herbivory accounted for greater than 20% of fine root biomass produced. Our study suggests that increases in sampling frequency from monthly to weekly can provide substantial gains in accuracy for estimates of root dynamics. Chapter 2 Abstract: We examined the impact of soil environmental variables (soil temperature, moisture, and available nitrate (NO3-) and ammonium (NH4+)) on the production, mortality, standing crop, turnover, and lifespan of Pinus palustris Miller fine roots using the minirhizotron technique. Data were collected for a full year from a P. palustris woodland in southwest Georgia. Mean soil temperatures appeared to have little influence on root processes, while temperature variance had a strong effect. More thermally variable microsites had increased root turnover and reduced root lifespans. Soil resources had a significant impact on demography; in particular, soil moisture and nitrate stimulated production, mortality, and turnover. High levels of soil resource availability also significantly reduced lifespan. Root lifespan was variable among individual roots based on root width, depth in the soil volume, and season of root production. Soil moisture had the strongest overall influence on root demography. This may result from the nature of our ecosystem (deep sands and subtropical climate); in addition, severe drought during our study may have enhanced the role of soil moisture, allowing environmental controls to increase in strength relative to within-plant controls on root demography. Master of Science

Understory light environments are inherently heterogeneous and therefore difficult to characterize. Numerous methods to measure understory light have been assessed in closed-canopied forests; however, the reliability of these methods has not been addressed for open-canopied forests. Therefore, the first objective of this study, presented in Chapter 3, was to test the accuracy and precision of various light measurement techniques at different time scales and sky conditions. The methods assessed performed differently depending on the sky condition and time of year when the sample was taken. To estimate annual photosynthetic photon flux density transmittance (annual %PPFD), the use of a 10-minute average of PPFD measured on an overcast day (%PPFDovercast) was effective, but accuracy decreased with decreasing solar altitude (ie season change). Hemispherical photographs used to estimate weighted canopy openness and gap fraction were effective methods, but gap light index (GLI) also derived from hemispherical photographs performed better. Accuracy of daily %PPFD estimates using %PPFDovercast, weighted canopy openness, and gap fraction were strongly affected by solar altitude and sky condition. Gap light index was very effective in estimating daily %PPFD for all sky conditions and time periods. The second objective of this study, presented in Chapter 4, was to characterize the relationship between canopy structure and spatial distribution of light by using three replicates of one uncut treatment and three harvest treatments: single tree, small gap (0.1 ha), and large gap (0.2 ha). Each harvest retained similar residual basal area but with different spatial patterns of the residuals, ranging from uniformly dispersed (single tree) to different degrees of aggregation (small and large gap). Average stand level light availability increased 12-22% when the same residual basal area of trees was distributed in clusters versus a uniform distribution. The variation of light availability increased as stands became more aggregated and larger amounts of the variation was explained by the spatial pattern of the canopy structure. Spatial autocorrelation range was twice as large in the small gap harvest then the other harvest treatments. It is suggested that seedling growth response to these differences in spatial patterns of light may differ between the different harvests. Master of Science

Spatial and temporal variation in foliar gas exchange on both a diurnal and seasonal scale was examined in 15-year-old fertilized and non-fertilized loblolly pine in the upper and lower thirds of crowns in stands located in the North Carolina sandhills. Photosynthesis rates between control and fertilized stands for both seasonal and diurnal measurement periods were different during only three months. Photosynthesis rates were consistently greater in the upper third of the crown compared to the lower third. Seasonal trends in both conductance and transpiration closely resembled trends found in seasonal photosynthesis. Foliar nitrogen concentrations were greater in fertilized stands for all months sampled. However, nitrogen content generally did not correlate with photosynthesis rates. Mean monthly water use efficiencies were significantly higher in fertilized stands during two months and were usually greater in upper crown foliage. Common empirical gas exchange models reveal that light and vapor pressure deficit (VPD) explain a majority of the variation observed in photosynthesis and transpiration, respectively. Conductance was not modeled since environmental variation did not adequately explain conductance patterns. Predicted light response curves reveal that upper crown foliage has higher maximum photosynthesis rates, respiration rates, light compensation points, and lower initial quantum yield compared to lower crown foliage. Models predict that foliage from fertilized stands is more sensitive to VPD and light during the growing season. Transpiration models predict highly variable responses to VPD depending on the treatment combination and season. Model R-square and predicted gas exchange values suggest that seasonal acclimation occurred. Master of Science

Purple-flowered rhododendron (Rhododendron ponticum L.) and yellow-flowered rhododendron (R. flavum Don.) are two dominant shrub species of the eastern beech (Fagus orientalis L.) understories in the eastern and western Black Sea Region (BSR), respectively. These invasive woody species significantly reduce beech growth and can preclude tree regeneration. The ecological consequence is an aging beech overstory with little or no regeneration to replace the mature trees. Great rhododendron (R. maximum L.) has been increasing in the forests of the Southern Appalachians of the United States, reducing tree regeneration and growth. The BSR and Southern Appalachians bear noteworthy similarities in climate, topography, and the forest flora. Purple-flowered and great rhododendrons also show important similarities in their ecology and the forest vegetation problems they can cause. Current rhododendron-dominated and threatened BSR forests may provide an advanced ecological picture of the forests of the Southern Appalachians in which great rhododendron now thrives. Therefore, new information gained on the ecology and effective and cost-efficient control of purple-flowered rhododendron may significantly improve forest management practices, not only for the current rhododendron-invaded BSR ecosystem, but also for other parts of the world. This dissertation consists of five separate yet related chapters. The first gives relevant literature reviewed for the dissertation. The second chapter focuses on various environmental and disturbance factors that may have shaped the current purple-flowered rhododendron-dominated beech forests of the BSR of Turkey. Chapter 3 assesses the effects of various manual and herbicidal woody control techniques on purple-flowered and yellow-flowered rhododendron in two field experiments in the BSR. The fourth chapter relates a study of uptake and translocation behavior of triclopyr ester and imazapyr in great rhododendron. This information is used to determine the optimum herbicide-surfactant combinations for the greatest active ingredient uptake and root translocation in great rhododendron. The last chapter is a synthesis of the information gained in all of these different experiments. Ph. D.

The Lower Coastal Plain of the southeastern United States is a major wood producing region. The region is characterized by a combination of nearly-level topography, poorly-drained soils, and high rainfall, which results in a perched water table in some soils that inundates the surface several times each year. Harvesting timber under wet site conditions often results in extensive soil compaction, rutting, soil displacement, and waterlogging. Forest managers are concerned that these visually-displeasing soil disturbances may cause site damage and reduced productivity. These concerns were addressed in an operational-scale field experiment conducted in South Carolina. The objectives of this experiment were to determine: (i) if soil disturbance changes key soil properties and processes; (ii) if soil disturbance reduces loblolly pine productivity; and, (iii) if disturbance can be mitigated with site preparation practices? Three 20-ha, 20-yr-old loblolly pine (Pinus taeda L.) plantations were harvested under wet and dry conditions to create a broad gradient in soil disturbance. Within each harvested plantation, a subset of 3-ha plots were site prepared by either bedding, or mole-plowing plus bedding, then all sites were established as 3rd -rotation pine plantations. Prior to site preparation, each plot was classified and mapped using a 5 by 5 soil disturbance (none to churned) by organic debris (none to slash piles) classification matrix. Within each plot, data were collected on several soil physical, chemical, and biological properties over a 2-yr period following site preparation. Key soil properties were integrated into a Soil Quality Index (SQI) and compared to aboveground productivity of 2-yr-old loblolly pine trees growing on closely-spaced (30 by 30 cm) bioassay plots planted across the gradient of soil disturbance. The soil physical properties were used to determine the least limiting water range (LLWR), the range in soil water content within which root growth is not limited. Soil compaction and deep rutting reduced the LLWR. Retention of logging slash improved the LLWR for compacted and rutted soils. Site preparation improved the quality of the soil physical environment across all levels of soil disturbance. Soil disturbance had no effect on soil chemical or biological properties as evidenced by no change in soil pH, ECEC, base saturation, available P, or net N mineralization with disturbance. The base saturation exceeded 80 % on all sites, with Ca saturation controlling soil pH. The high base saturation buffered any redox-induced changes in soil chemistry that would have resulted from disturbance. The results showed that high fertility is an important mechanism for buffering the potentially-negative effects of soil disturbance on the soil nutritional environment. Site preparation changed soil chemical properties, but the changes were probably associated with tillage effects on organic matter and clay content, not redox processes. The SQI showed that surface soil compaction and deep rutting reduced soil quality, mainly by decreasing the LLWR and aeration depth. Site preparation mitigated the effects of most disturbances on soil quality, evidenced by similar aboveground biomass production among soil disturbance classes after bedding. A regression model was developed for predicting aboveground biomass production as a function of SQI. SQI explained 73 % of the variation in aboveground biomass production. The regression model showed that compression tracks and rutting decreased aboveground biomass production compared to undisturbed soils. The long-term effect of these disturbances on productivity will depend on natural soil recovery processes. However, these early results suggest that compaction and rutting should be minimized on similar sites, especially if sites will not be bedded before reforestation. The mole-plow / bedding treatment increased aboveground biomass production, indicating that this experimental treatment may be a viable practice for enhancing productivity. Ph. D.

A study was conducted at the Savannah River Site near Aiken, SC to: 1) characterize the spatial patterns of soil and forest floor variables (moisture, pH, soil phosphate, forest floor and soil carbon and nitrogen, and soil available nitrogen), 2) assess the spatial patterns of the plant community, and 3) investigate spatial relationships among the variables and between the variables and woody vegetation. Spatial soil and litter samples were collected on five 0.25 hectare plots, and relationships were explored using Pearson's correlation tests, canonical correlation analysis, variogram modeling and kriging. The average range of spatial autocorrelation for the forest floor variables was >45 m, while that for soil variables was 12 m. Woody stem basal area exhibited spatial autocorrelation at ranges of less than 12 m, and was only weakly correlated with forest floor and soil resource patterns. Few strong spatial correlations among the forest floor and soil variables were observed. The means and variances of the variables were low, and differences in resource levels probably had little impact on the spatial pattern of vegetation. Results indicate a weak, differential effect of species group on litter quality, a weak relationship between large pine trees and soil nitrogen patterns, and a general homogeneity of the stands. Master of Science

Effects of thinning, such as increased diameter growth and decreased mortality in the residual stand, have been recognized by foresters for many years. These effects are largely the result of increased tree vigor which is induced by a decreased level of competition. These relationships are reflected in many of the models that are central to PTAEDA2, a growth and yield simulator which was developed for use with loblolly pine (Pinus taeda) plantations established on cut-over, site prepared lands. Data from a long-term thinning study served as a basis for attempting to improve the predictive output of PTAEDA2. Assessment of differences in model parameter estimates between three levels of thinning intensity led to various approaches to reach this goal. Height increment and mortality models were found to need no additional refinement and were re-fit using all available data. Diameter increment and crown ratio model forms could not account for thinning effects in their present form and thinning response functions that could provide the proper behavioral response were added to these models.Models were evaluated individually and in combinations in a reduced growth simulator. This reduced simulator is a modified form of the growth subroutines in PTAEDA2 and is designed to utilize external data. Results of growth simulation runs show improvements in predictive ability for the crown ratio model fit to all data and for the re-fit height increment model/crown ratio model with thinning response variable combination. The diameter increment model with a thinning response variable significantly improved diameter prediction within the simulator, but predicted stand volumes were poor. The re-fit mortality function resulted in greater prediction error for mortality than the original PTAEDA2 mortality function. Master of Science

Plant root system response to nutrient heterogeneity was tested in ten plant species of varying life form and successional status. All plants tested are native to the South Carolina coastal plain. Morphological responses of the root system (scale, precision and discrimination) and overall plant response (sensitivity) to increasing nutrient heterogeneity were tested. Ten individuals of each species were placed into four treatments which had varying nutrient distribution but the same overall nutrient addition. Plants were harvested when roots reached pot edge. I observed high variation in scale (mass and extent of a root system), precision (the ability to proliferate roots in nutrient patches) and sensitivity (growth benefits gained as nutrient heterogeneity increases; measured as total biomass). No significant discrimination responses were observed, although greatest mean root density occurred at intermediate fertility levels for all species. I tested the hypothesis that scale and precision would be negatively correlated, and I did not observe this relationship in these plant species. However, in herbaceous species scale and precision were positively correlated. Sensitivity was not closely related to precision indicating that proliferating roots in fertile patches does not always yield growth benefits in heterogeneous soils. Further, some sensitive species had very low precision suggesting that other characteristics lead to positive growth response in heterogeneous environments. Plasticity of root uptake rates and demography of roots are proposed as two other mechanisms which may play important roles in plant sensitivity responses. Scale was negatively correlated to sensitivity for herbaceous plants suggesting that plants that monopolize the most soil space are not able to gain benefits from nutrient patches within the soil matrix. There was no trend observed to suggest that plant life form was correlated with precision or sensitivity. However, scale was greater in herbs than in woody plants, possibly because the two life forms develop at different times. Master of Science

Use of soil amendments at planting is one of the time-honored traditions in horticulture, although their effectiveness has been questioned by many. Recently, humate and humate-based products, generally known as biostimulants, have been marketed to increase transplant success. In this study, three experiments were conducted to examine the effects of soil amendments and biostimulants on post-transplant growth of landscape trees. The first experiment, conducted in a greenhouse, determined the effects of several biostimulant treatments (granular humate, water-soluble humate, liquid humate, liquid humate+ = humic acid, hormones, and vitamins) and fertilizer levels (low, medium, high) on the growth of container-grown Corylus colurna L. (Turkish hazelnut) seedlings. Biostimulants did not increase top growth compared to control treatments, but root growth was increased by granular humate at a medium fertilizer rate. The second experiment examined the effects of biostimulants (granular humate, water-soluble humate, liquid humate+) on the post-transplant root growth and sap-flow of landscape-sized balled and burlapped Acer rubrum L. (red maple) grown in root observation compartments (rhizotrons). Biostimulants did not increase root growth over control treatments, but sap-flow was increased. The third experiment, conducted in the field (Groseclose silt loam soil) investigated the effects of soil amendments (peat, and compost) and biostimulants (granular humate, and liquid humate+) on the post-transplant growth of Crataegus phaenopyrum (Blume) Hara (Washington hawthorn) and red maple transplanted bare-root, and grown under combinations of irrigated vs non-irrigated and fertilized-at-planting vs non-fertilized-at-planting regimes. Hawthorn controls generally had less top growth than the other soil treatments as a whole. No soil treatment was higher than control for top growth of red maple. However, root growth of red maple was highest in the peat-treated trees. Stem diameter and dry mass for the control and compost treatments were higher than the biostimulant treatments in irrigated plots, but no differences were observed in non-irrigated plots. Granular humate-treated trees resulted in higher stem diameter and dry mass than the liquid humate+-treated trees in non-irrigated plots. There were no effects of fertilizer, or irrigation on growth after two growing seasons for either species. Master of Science

Nine provenances of Turkish red pine (Pinus brutia Ten.), an eastern Mediterranean conifer, taken from a wide range of locations in Turkey, were grown in individual pots in a greenhouse either with or without joint goose goatgrass (Aegilops cylindrica L.) under either high or low soil water availability for two growing seasons. The study consisted of two separate experiments started at times with varying microenvironmental conditions and Turkish red pine (TRP) provenances. Diameter and secondary needle, stem, and root biomass were measured at the end of the second growing season, while height was measured after both the first and second growing seasons. Pine stem water potential (SWP), measured at the end of the second growing season, was the only physiological variable in the study. Master of Science