For this dissertation, I measured how plant communities in Appalachian forests responded to disturbances caused by forest management activities. I had two primary objectives including (1) testing theories of biological diversity and invasions by nonnative species; and (2) providing empirical data that will help guide the sustainable use of forest resources. This work is part of the Southern Appalachian Silviculture and Biodiversity (SASAB) experiment that was established in the early 1990s to investigate ecosystem responses to a gradient of timber harvesting disturbances. Ranging from undisturbed controls to silvicultural clearcuts, the disturbance gradient is replicated at sites located throughout the Appalachian mountains of Virginia and West Virginia. The plant community was sampled across a wide range of spatial scales (2 hectares to 1 m2) using a nested sampling design and was also sampled at three times including pre-disturbance, one year post-disturbance, and ten year post-disturbance. For one element of the study I tested modern theories of biological invasions and investigated how the forest disturbance gradient interacted with species diversity to influence invasion by nonnative species (Chapter 2). Contrary to popular ecological theories of biotic resistance, the most diverse sites tended to be more easily invaded following intense canopy disturbance. Interestingly, none of the sites in this study were dominated by invasive plants, which led me to ask whether dominant tree species in forests provided resistance to nonnative plant establishment and growth through the quality of the litter they produce. I also asked how might animals that are known to alter litter layers interact with dominant tree species to influence plant invasions. Therefore, I conducted an experiment investigating how changes in litter from dominant tree species and invasions by nonnative earthworms might influence invasibility of forests using forest floor mesocosms (Chapter 3). I found that plant invasion was inhibited by native oak litter even when earthworms were present, suggesting that oak forests may resist plant invasions via oak tree litter. In contrast, plant invasion was greater under invasive tree litter and earthworm activity tended to facilitate invasive plant success only under invasive tree litter. I was also interested in understanding how disturbance might alter relationships between local and regional diversity. The long-term data of the forest disturbance experiment allowed me to investigate how local species richness is mediated by regional species richness after disturbance and during forest community development (Chapter 4). Local richness depended strongly on regional richness only after disturbance via colonization of species, but this relationship changed during forest aggradation. These results suggest that regional species pools are important to maintain local diversity following disturbance, but that local interactions (through canopy closure of dominant trees) exert control over species diversity during community reorganization. Lastly, I tested current theories on how diversity influences compositional stability after disturbance (Chapter 5). Disturbance consistently resulted in decreased compositional stability, but diversity was associated with stability in complex ways, which depended on how stability was measured and at what scale. Species-rich areas were in some instances less stable; in other instances areas with intermediate levels of diversity were more stable. These results suggest that disturbance causes shifts in species composition via colonization, but the ways in which diversity of sites influences compositional stability is complex and depends on methods used and the scales of observation. Taken together, these results suggest that disturbance influences invasibility, species saturation, and compositional stability of ecological communities. These properties change immediately following disturbance, and during forest development and canopy closure. Data from this project were useful in testing existing theories of community ecology, and may ultimately prove useful for forest managers as they decide how to protect biodiversity while planning for other uses of forest resources. Overall, these results suggest that colonization of species is the primary process driving plant community patterns in Appalachian forests following disturbance. / Ph. D.

The passage of the Federal Water Pollution Control Act of 1972 required states to develop forestry BMPs to help reduce potential nonpoint source pollution from forest operations. Properly applied forestry best management practices (BMPs) have since been proven to protect water quality from forest operations. This research project reviewed BMP effectiveness studies in the U.S., assessed current state developed and implemented of forestry BMPs, and developed a simple method to estimate potential erosion from forest operations for the Piedmont physiographic region based on previous studies. Eighty-one BMP effectiveness studies were reviewed. The review of past effectiveness studies indicates that water quality protection is increased when BMPs are implemented correctly. These effectiveness studies provide states with valuable information on how their BMP guidelines are achieving the goals defined by the Federal Water Pollution Control Act. Every U.S. state has forestry BMP guidelines. These guidelines may be non-regulatory, quasi-regulatory, or regulatory depending on the state. Twenty states reported implementing non-regulatory BMP guidelines, 19 quasi-regulatory BMP guidelines, and 11 regulatory BMP guidelines. State forestry agencies were reported as being the lead agency responsible for BMP monitoring in 35 states. The national forestry BMP implementation rate was 91% (32 states). However, states did report deficiencies for specific BMP guideline categories. Supplementary to the reviewed BMP effectiveness studies, forest erosion studies in the southeastern U.S. that quantified erosion rates from forest operations were also reviewed. Erosion rates obtained from the literature were reviewed by operation categories (timber harvesting, forest roads, skid trails, log landings, stream crossings, and streamside management zones) and physiographic region (Mountains, Piedmont, Gulf Coastal Plain, and Atlantic Coastal Plain). There were numerous research gaps regarding erosion rates from forest operations for all the regions except the Piedmont region. The Piedmont region was selected for developing a method to estimate potential erosion from forest operations. This erosion estimation method is a quick and potentially useful tool for estimating potential erosion; however, it is based on limited data from the Piedmont region only. The basic method approach might be considered for the other physiographic regions, but further research is needed to fill current knowledge gaps. / Ph. D.

Interest in wood-based bio-energy systems in the United States is increasing and may play a part in future renewable energy initiatives (Dincer 2000). Family forests have potential to play an important role in supplying wood biomass for energy production. However, access depends mostly on the management intentions among family forest owners. Enhanced biomass markets in regions where family forest ownership dominates could increase productivity by reinvigorating the low-value merchandizing required to accomplish silvicultural objectives. Given diverse owner objectives and forest types on family forests, estimates of biomass availability must include both biophysical and social aspects of procurable feedstock. This thesis chronicles a biosocial case study that estimates potential biomass supply from 51 family forests in Virginia and North Carolina. The study occurred within a woodshed centered on the future site of an impending ethanol plant in Mecklenburg County, Virginia. A survey instrument using the theory of planned behavior was used to measure ownership characteristics and intention to harvest. Forest attributes were collected during property visits to estimate potential yields resulting from silvicultural simulations. Results reveal that forest cover-type and tree size significantly affect owner intentions to harvest and owner attitudes toward harvesting partially mediate this relationship. Outputs from silvicultural simulations correspond with those made using Forest Inventory and Analysis data within the study region. Disproportionality was examined by coupling social and biological drivers of sustainable wood biomass availability. Implications of the research include refined estimates of potential supply and demonstrating a multi-scalar, mixed-method approach for assessing wood biomass availability. / Master of Science

Solitary Vireos (Vireo solitarius alticola) were studied on an extensive mature forest landscape in 1994 and on three sites harvested using the group selection method in 1995. Pairing success was 88% on the unharvested site in 1994, however it was lower on the harvested sites in 1995 (53.8%). Low reproductive success was observed in both years (12% and 22% respectively). In 1995, the amount of opening within a 200 meter radius of the first observation point of a male was positively related to pairing success (p<0.05). This suggests that females may be cueing in on the increased number of shrubs in and around openings. / Master of Science

Amphibians, because of their semi-permeable skin, sensitivity to changing microclimates, and important role in ecosystems, are often viewed as indicators of ecosystem health. They make excellent organisms for studies on the effects of silvicultural practices. My goal was to provide recommendations for forest management in the southern Appalachians so that harvesting operations are compatible with maintaining healthy populations of forest amphibians. I tested previously created habitat models that determined the most important habitat characteristics for salamanders. I counted salamanders in 240 10 x 10 m plots located in the MeadWestvaco Wildlife and Ecosystem Research Forest in north-central West Virginia. We also collected a variety of habitat data in these plots to predict salamander abundance with previously created models. These simple linear regression analyses of predicted versus observed values suggest for most models (7 out of 9) a weak relationship between predicted and observed values (R2 from 0.0033 to 0.2869, p from < 0.0001 to 0.7490). However, one of the models showed characteristics suggesting that it predicted new data as well or better than the original data, and therefore was the most accurate at predicting salamander abundance, and could be used for management purposes, although there was still much unexplained variation. This model included the variables woody stems (< 7.5 cm DBH), available rock, riparian status (i.e., within 15 m of a stream), percent overstory canopy cover, and available highly decomposed woody debris (decomposition classes 3 to 5). All of these relationships were positive except for woody stems, suggesting that in order to maintain healthy populations of salamanders, we should protect areas next to streams, with high amounts of rock, decomposed woody debris, overstory canopy cover, and few woody stems. I also examined the immediate effects of clearcuts on salamander movement and population estimates. I batch marked salamanders in plots at the edges of a clearcut, and in a control plot. Using the Schnabel estimator, I estimated population sizes in each plot. I then compared population estimates pre- and post-harvest on the interior (harvested) and exterior (unharvested) sides of the plots, taking into account the control plot. I also examined adult-juvenile ratios and movements from one side of the plot to the other. I found no significant changes (p > 0.05) following harvest in any of these measures, suggesting that salamanders do not move out of the harvested area post-harvest, at least over the short term (10 months of this study). This suggests that a longer period of time (> 1 year) is required to observe the population declines detected in most studies. / Master of Science

Varietal forestry may increase the productivity of loblolly pine (Pinus taeda L.) in the Southern United Sates. However, the effects of these genetic x environment interactions are still poorly understood. In this study we examined the responses in growth, stand uniformity and leaf level physiology of loblolly pine clonal varieties and families to silvicultural intensity and site effects. We also looked for patterns in observed traits that were consistent between crown ideotypes. Two varieties of each crown ideotype (narrow vs broad crowns) and two families (controlled mass pollinated (CMP) and open pollinated (OP) family) were tested on the Virginia Piedmont (VA) and the North Carolina Coastal Plain (NC) under different silvicultural intensities (operational vs intensive), and planting density (617, 1235 and 1852 trees per hectare). Data were collected during the first four growing season after establishment. At NC, intensive silviculture increased crown-width, height and dbh by 33%, 14%, and 23%, respectively. At VA, intensive silviculture increased crown-width, height and dbh by 41%, 10%, and 23%, respectively. Intensive silviculture also increased slightly but significantly the stand uniformity of stem growth. However, the differences in productivity between silvicultural treatments were not explained by differences in leaf-level physiology. Across all treatments and sites the varieties generally grew faster than the OP family, but the differences were higher at VA. Varieties did not differ in stem growth, but the broadest crown variety had greater stand uniformity, photosynthetic rate (Asat), carbon isotope discrimination (∆¹³) and lower fascicle size than the OP family. None of the traits assessed inthis study was consistent within the ideotypes. Varieties classified in the same crown-ideotypes may respond differently to the environmental effects of site and silviculture, which reinforces the need of matching varietal forestry with precision silviculture to achieve gains in productivity. / Ph. D.

Over one million hectares of pocosins and wet flats in the southeastern coastal plain are intensively managed for the production of loblolly pine (Pinus taeda) plantations. These management activities may have adverse effects on soil physical properties, site hydrology, and overall site productivity. Substantial quantities of wood residues are often left on these sites by timber harvesting operations, and it was hypothesized that the incorporation of this slash into the soil could improve the soil physical properties and site hydrology. One organic pocosin site and one mineral wet flat site were chosen post-harvest for treatment. The wet flat study was organized as an incomplete block design having four blocks and six treatments: (i) conventional bedding, (ii) strip surface mulching with bedding, (iii) strip surface mulching with tillage and bedding, (iv) broadcast mulch without bedding, (v) broadcast mulch with bedding, and (vi) flat planted control. The pocosin study was organized as a randomized complete block design with four blocks and four treatments. The treatments are identical to those of the wet flat site without the broadcast mulch treatments (iv and v). Soil physical property data was analyzed pre- and post-treatment, while post-treatment site hydrology and soil water chemistry data was analyzed periodically for one year. Seedling survival and height data were analyzed after one growing season. The treatments had little effect on soil physical properties, site hydrology, soil water nutrients, or seedling survival on the wet flat study site. Bedding in general significantly increased tree height growth, but mulching had no significant effects. The treatments had little effect on soil physical properties on the pocosin study site except for soil macroporosity, which was significantly increased by bedding. Site hydrology and soil water nutrients were not significantly affected by the treatments, but seedling survival and height growth were significantly increased by bedding. Mulching had no significant effects on any of the parameters studied. / Master of Science

Harvesting practices in the southern Appalachians have moved away from clearcutting in favor of variable retention harvesting systems. A study was initiated in 1995-8 to investigate the effects of retaining varying numbers of residual trees on regeneration in seven silvicultural treatments. A second study specifically focused on stump sprouting in only three of those treatments. The treatments for first study included: a clearcut, commercial harvest, leave-tree, shelterwood, group selection, midstory treatment, and an uncut control. The second only focused on the clearcut, leave-tree, and shelterwood. These treatments were implemented in seven stands in Virginia and West Virginia over two physiographic provinces, the Appalachian plateau and Ridge and Valley. The stands were even-aged oak dominated Appalachian hardwood stands on fair quality sites with average ages ranging from 63 to 100 yrs. Permanent plots were randomly located in each stand and all overstory trees (>5m tall) were inventoried and tagged prior to harvest. Regeneration was also quantified. Harvest occurred between 1995-8. For the current studies the plots were re-inventoried 9-11 years post-harvest and all regeneration in all treatments as well as stump sprouts in the selected treatments were quantified. The first study utilized a mixed model ANOVA to analyze five species groups: oak, maple, black cherry-yellow-poplar, miscellaneous, and midstory. Response variables included importance value, average height, and density compared within species group and among treatments. Differences between sprout and seedling origin regeneration were also investigated within species group among treatment. Results indicated that oak densities were similar in all of the treatments, and stump sprouts were larger and more frequent than seedlings. Maple exhibited an increase from pre-harvest overstory importance and exhibited competitive sprouting. The black cherry-yellow-poplar group had few but highly competitive sprouts and a considerable increase in seedling origin regeneration in all treatments. The miscellaneous species densities increased as well with more competitive sprouting in some treatments. The midstory species were excluded from the analysis as it was assumed these species would not occupy canopy positions in a mature stand. The second study investigated differences in the percent of stumps that sprouted and the number of sprouts per stump. The percent data were analyzed using a non-parametric one-way ANOVA and regression analysis, while the sprouts per stump data were compared in a mixed model ANOVA and regression. Species were combined into six groups: the red oak group, chestnut oak, red maple, white oak/hickory group, mixed mesic group, and midstory group. The plateau tended to have reduced sprouting compared to the Ridge and Valley for most species groups and treatments. The red oak group, chestnut oak, and red maple exhibited reduced sprouting with increased residual basal area. The mixed mesic group did not show any effect in sprouting related to residual basal area. Only chestnut oak showed fewer sprouts per stump as residual basal area increased. / Master of Science

Chemical site preparation is used to enhance the productivity of loblolly pine plantations; however, it is most often combined with other methods and/or chemical release, and has been studied little on its own. Our study, conducted in the Virginia piedmont, compares the effects of various site preparation applications of imazapyr and their timings (July 23rd, September 3rd, and October 1st 2005) against a year-two chemical release (September 12th 2007), and an untreated check, all following a site preparation burn (June 15, 2005). Testing for additive effects, site preparation treatments were conducted with and without, sulfometuron methyl (SMAX) which targets herbaceous vegetation. Half of all treatments, including the check and release, received a year-one weed control treatment (H) of imazapyr and SMAX. Pines were measured every few years, including the latest measurement in year 14 (2019). Competing hardwood vegetation basal area was measured in year 14. All site preparation treatments reduced competition levels compared to the check, and an inverse relationship exists between competition level and pine volume. Adding SMAX resulted in less effective competition control, but had little effect on pine volumes. Adding H decreased average competition levels, and increased pine volumes. H treatments had the most effect at improving early season (July) applications. September and October treatments generally had more effect than July and the year-two release at controlling competing vegetation, and also showed higher individual pine tree volumes than all other treatments. Chemical site preparation treatments, even performed alone, appear to be valid means of improving pine stands. / Master of Science / Competing vegetation has a negative effect on planted pine production. Areas are often prepared with herbicides or mechanical removal of vegetation before crop-trees are planted (site preparation), or receive herbicide treatments after planting (release), to reduce levels of competition. Our study focuses on pre-planting herbicide (imazapyr with and without sulfometuron methyl) use at different timings (July 23rd, September 3rd, and October 1st). A chemical that targets herbaceous vegetation (sulfometuron methyl) was included in half of the pre-planting treatments, and half of all treatments, including the otherwise untreated check and year-two release, received an added weed control treatment in year-one. This was done to test for potential additive effects of chemical vegetation control options on pine productivity. September and October treatments have a greater effect at controlling competition 14 years since treatment and increasing pine volumes than the July, year-two release, and check. The added herbaceous control increased competition levels on average, without having an effect on the planted pine volumes. The year-one weed control decreased competition levels, and increased pine productivity, particularly within the July treatments. Overall, preparing sites with herbicide is beneficial to the pine stand, even when performed alone. Adding the herbaceous vegetation control is likely not necessary, and the extra year-one weed control may provide a boost to herbicide treatments applied earlier in the season.

Survival and growth of planted trees are common indices used to evaluate success of wetland restoration efforts used to compensate for wetland losses. Restoration efforts on marginal agricultural lands typically result in less than satisfactory survival and growth of desired tree species. This study evaluated seed source ecotype, greenhouse preconditioning and combinations of five mechanical site preparation techniques (mound, bed, rip, disk, pit), four levels of planting stock (gallon, tubeling, bare root, and direct seed), and three planting aids (mat, tube, none) on the survival and growth of American sycamore (Platanus occidentalis L.) and willow oak (Quercus phellos L.) planted on an old field riparian area in the Virginia Piedmont. American sycamore seedlings subjected to greenhouse flood preconditioning had 25% greater height and willow oak seedlings grown under normal greenhouse conditions had 18% greater diameter, but these greenhouse adaptations did not confer greater survivability or growth after field planting. American sycamore seeds sourced from dry ecotypes were 14% taller than wet ecotype seeds, and willow oak acorns sourced from wet ecotypes were 11% taller than dry ecotype acorns, indicating that parental ecotype may influence survivability and growth. The combination of mounding site preparation and gallon planting stock increased mean survival to 100% and aboveground dry biomass (5.44 Mg/ha/yr) in American sycamore. Willow oak had 45% greater woody stem volume with mounding site preparation 80% greater woody stem volume with gallon and bare root planting stock. Tubeling planting stock provided significant benefit relative to the low planting stock cost for American sycamore, while bare root seedlings were shown to be an effective planting stock for willow oak. The use of appropriate ecotype seed sources, use of mounding mechanical site preparation techniques and planting of species appropriate planting stock increased survival and growth of common early and mid-successional Piedmont tree species on marginal agricultural lands. Treatments that appear to be economically viable for restoration and mitigation efforts could potentially offer other economic incentives such as short rotation woody crops and timber value, which might induce additional private landowners to attempt restoration efforts in marginal old field riparian areas. / Doctor of Philosophy / In order to offset the loss of wetlands due to development, strategies are needed to create wetlands in areas along streams that are unfarmed. Survival and growth of planted trees are commonly used to evaluate the success of these new wetland areas. The goal of this research is to provide alternative methods to increase survival and growth of two common trees planted in the Piedmont of Virginia. The results show that creating mounds of soil before planting trees and planting larger trees will increase tree survival and growth in these wetland areas. Landowners and land managers can use these methods to increase the value of unfarmed land along streams while also increasing water quality and providing habitat animals that use the streams and wetlands.