Suppression of canopy tree recruitment beneath rapidly spreading thickets of Rhododendron maximum L. (Ericaceae) in southern Appalachian forests is an issue of major concern because of the potential impacts on forest productivity, hydrology and wildlife habitat. Many studies have investigated the causes of seedling inhibition beneath dense shrub understories, but few have uncovered specific mechanisms leading to seedling decline. In this study, I have examined the influence of the evergreen understory (R. maximum and Kalmia latifolia L.) on tree recruitment processes at multiple stages - seed rain, seed bank, and post-establishment seedling growth and survivorship. Effects of dense shrub cover on seed rain and seed bank density and composition were examined using a paired treatment design in which samples were collected beneath shrub-influenced and open understories. A second experiment investigated the influence of R. maximum and K. latifolia density on the growth and survivorship of Quercus seedlings, resource availability, and the rates / causes of seedling damage. I found that neither seed rain, nor seed bank density or species richness was inhibited by the presence of R. maximum or K. latifolia. Forest seed banks were dominated by sweet birch (Betula lenta L.), and were compositionally disparate from the overstory. Analysis of resource competition between shrubs and seedlings indicated that seedling performance and survivorship was a negative function of R. maximum density. Open-canopy light availability, nitrogen content in the organic horizon (litter and humus), and soil nutrient availability were potential resource-related mechanisms. Further, I found that the rates of insect herbivory on Quercus seedlings were positively correlated with R. maximum density. Kalmia latifolia had little influence on resource availability, seedling performance or herbivory rates, and does not appear to have a suppressive effect on tree seedlings. Overall, this research indicates that resource competition is the primary mechanism by which seedling suppression occurs beneath R. maximum, and that increased herbivory on seedlings may be an additional mechanism that demands further study. / Master of Science

<I>Rhododendron maximum</I> L. (Ericaceae) is an evergreen shrub that grows in dense thickets and currently covers large areas of the understory in the deciduous forests of the southeastern United States. Thickets of R. maximum are inhibitory to recruitment and regeneration of many understory plants including canopy tree seedlings. By effectively lowering the survivorship of woody species trying to establish within thickets, <I>R. maximum</I> could influence stand level regeneration patterns and ultimately the community structure of these deciduous forests. This dissertation outlines research conducted to determine if: 1) below and above ground resources are lower within thickets of <I>R. maximum</I> when compared to forest sites where <I>R. maximum</I> is absent; 2) <I>Quercus rubra</I> and <I>Prunus serotina</I> seedlings growing in thickets have lower mid-day photosynthetic rates; 3) <I>Quercus rubra</I> and <I>Prunus serotina</I> seedlings growing within thickets are low light acclimated when compared to seedlings growing in forest without <I>R. maximum</I>; 4) the presence of <I>R. maximum</I> constrains CO₂ assimilation of <I>Quercus rubra</I> seedlings exposed to light flecks of different durations and intensities; 5) the presence of <I>R. maximum</I> constrains the light fleck responses of <I>Quercus rubra</I> seedlings exposed to eight light flecks in rapid succession; and 6) canopy openness regulates the capacity of <I>Quercus rubra</I> seedlings to assimilate carbon when exposed to eight consecutive light flecks. <I>Rhododendron maximum</I> thickets altered resource availability for seedlings when compared to areas of forest without <I>R. maximum</I>. Diffused photosynthetically active radiation (PAR) averaged less than 5 μmol m⁻² s⁻¹ throughout the growing season in sites with <I>R. maximum</I> in comparison to 10-30 μmol m⁻² s⁻¹ in sites without <I>R. maximum</I>. Soil moisture content, measured using Time Domain Reflectometry was approximately 6% lower in forest sites with <I>R. maximum</I> compared to sites without <I>R. maximum</I> throughout the growing season. Most nutrient concentrations (e.g.,, C, N and most cations) and nitrogen mineralization rates were significantly lower in sites with <I>R. maximum</I>. Temperature and atmospheric relative humidity are slightly lower under thickets of <I>R. maximum</I>. In general, sites with <I>R. maximum</I> are associated with lower resource availability above and below ground in comparison with sites without <I>R. maximum</I>. Attenuation of below canopy PAR by thickets of <I>R. maximum</I> negatively influences the photosynthetic capacity of <I>Quercus rubra</I> and <I>Prunus serotina</I> seedlings as indicated by measurements of mid-day photosynthesis. In 1996, the seasonal mean mid-day photosynthetic rate of first year <I>Q. rubra</I> seedlings growing in <I>R. maximum</I> thickets (1.3 μmol m⁻² s⁻¹) was 62% lower than the seasonal mean mid-day photosynthetic rate (2.1 μmol m⁻² s⁻¹) of seedlings growing in forest sites without <I>R. maximum</I>. For second year seedlings in 1997, seasonal mean mid-day photosynthesis was 183% higher for plants growing outside of thickets (1.7 μmol m⁻² s⁻¹) compared to the mean rate (0.6 μmol m⁻² s⁻¹) for plants located within thicket sites. The mean mid-day PAR available to seedlings located in forest sites without <I>R. maximum</I> during measurements of photosynthesis was 354% higher in 1996 and 257% higher in 1997. First year <I>Prunus serotina</I> seedlings growing in forest without <I>R. maximum</I> also had greater seasonal mean mid-day photosynthesis (0.7 μmol m⁻² s⁻¹) when compared to the mean rate (-0.1 μmol m⁻² s⁻¹) for plants growing within thickets. <I>Prunus serotina</I> seedlings located in the presence of <I>R. maximum</I> received on average 67% less PAR. Photosynthetic acclimation to low light was assessed for <I>Q. rubra</I> and <I>P. serotina</I> seedlings growing under both forest conditions by measuring photosynthetic responses to light <I>in situ</I> using even aged one-year old seedlings. <I>Quercus rubra</I> seedlings growing in forest sites without <I>R. maximum</I> had significantly higher light saturated rates of photosynthesis. For both species, photosynthetic responses to light were otherwise similar irrespective of the presence or absence of <I>R. maximum</I>. The impact of the <I>R. maximum</I> subcanopy on understory PAR and subsequent influence on canopy tree seedling photosynthetic capacity implies that sunflecks are critical for seedling net carbon gain in these forest understory environments. To determine the effect of <I>R. maximum</I> on the photosynthetic response to sunflecks of oak seedlings, light flecks were simulated on 288 randomly chosen, even aged, two-year old seedlings <I>in situ</I>. Half of the seedlings were located within <I>R. maximum</I> thickets. Seedlings were randomly assigned one of four light fleck durations (30, 60, 120, and 300s) and one of three intensities (100, 500, 1000 μmol m⁻² s⁻¹). Half of all seedlings were dark pre-acclimated prior to light fleck simulations by covering with aluminum foil for at least 12 hours, while the remaining seedlings were pre-acclimated under ambient conditions. Analysis of covariance showed that a significant, positive, linear relationship exists between the length of a light fleck and total carbon gain during a light fleck for seedlings in forest sites with and without <I>R. maximum</I> regardless of pre-acclimation status, or light fleck intensity. Furthermore, there was a significant effect of <I>R. maximum</I> on the slope of the relationship such that following ambient pre-acclimation, seedlings located within thickets assimilated significantly less carbon with increasing light fleck length than seedlings located in forest sites without <I>R. maximum</I>. When seedlings were dark pre-acclimated there was no difference in carbon gain with increasing fleck length between seedlings in forest with and without <I>R. maximum</I> except for flecks of 1000 μmol m⁻² s⁻¹. The data lead to the conclusion that under natural conditions the presence of <I>R. maximum</I> likely prohibits <I>Q. rubra</I> seedlings from utilizing sunflecks as effectively as seedlings growing in forest sites where <I>R. maximum</I> is absent. Because sunflecks often occur clustered together during a short period of time during the day, another field study was conducted to further characterize the effect of <I>R. maximum</I> on the photosynthetic response of oak seedlings to eight consecutive light flecks. Within 10 paired sites, (i.e., with and without <I>R. maximum</I>) 3 even aged three-year old <I>Q. rubra</I> seedlings were selected. Over each seedling, a hemispherical canopy photograph was taken and analyzed for percent canopy openness. Each seedling was dark pre-acclimated for 12 hours and then exposed to eight light flecks in rapid succession during which time photosynthesis was logged every two seconds. Each light fleck was 500 μmol m⁻² s⁻¹ in intensity and lasted for 120s. Following each light fleck, leaves were exposed to 10 μmol m⁻² s⁻¹ PAR for 60s before the next light fleck. Mean carbon gain and maximum photosynthesis achieved during each light fleck was significantly lower for seedlings located in the presence of <I>R. maximum</I> for all flecks in an eight-fleck simulation. In addition, seedlings located within thickets generally had significantly lower pre-illumination photosynthesis following the first of eight light flecks. The mean photosynthetic light use efficiency of seedlings located in forest with <I>R. maximum</I> was significantly lower for the first six of eight light flecks in succession. Using regression analysis and analysis of covariance, percent canopy openness was used to explain the variation in carbon gained from all eight light flecks in succession for seedlings under both forest conditions. However, significant relationships failed to exist between under either forest condition and precluded using analysis of covariance. The results from these studies lead to the conclusion that light limitation is a major mechanism responsible for the extirpation of canopy tree seedlings from within thickets of <I>R. maximum</I>. Tree seedlings growing in forest sites with <I>R. maximum</I> receive less solar irradiance, have lower mid-day photosynthesis, fail to acclimate to the lower light conditions within thickets, and utilize sunflecks less effectively as well as less efficiently when compared to plants growing in forest sites without <I>R. maximum</I>. / Ph. D.

Rhododendron maximum L. is a dominant subcanopy species in the southern Appalachian Mountains. R. maximum undergo distinct thermonastic leaf movements (TLM). The purpose of these movements has not yet been determined. Previous studies have suggested TLM are a photoprotective mechanism for the dynamic light environment of the subcanopy in a deciduous forest during winter. The present study aimed to determine the effects of restricting TLM on photoinhibition, net photosynthesis, and other gas exchange parameters, particularly during the early spring. After restricting TLM on certain leaves, we observed the above parameters from autumn 2005 to late spring 2006. Our results indicated that photoinhibition increased (lower Fv/Fm) in treatment leaves over reference leaves throughout the winter. The difference became greater during the early spring, when reference leaves began to return to normal levels of photochemical efficiency and treatment leaves sustained low Fv/Fm. Net photosynthesis was lower for treatment leaves than reference leaves. This became most significant during the early spring, when maximum carbon gain is possible. Finally, gas exchange parameters as measured by light and CO2 response curves did not indicate any significant difference between treatment and reference leaves post canopy closure. Out results suggest that TLM are an important mechanism for photoprotection, allowing leaves of R. maximum to recover quickly during the early spring and maximize their early spring carbon gain. / Master of Science

The literature was reviewed for information regarding anatomical studies of secondary xyqlem - particularly information pertaining to <u>Acer Nigrum, Platanus occidentalis</u>, and <u>Rhododendron maximum</u>. / Master of Science

Rosebay rhododendron (Rhododendron maximum L.) is an evergreen ericaceous shrub that plays a dynamic role in the southern Appalachian forests. Commonly located on mesic sites, this understory shrub forms dense thickets that greatly reduce the amount of light available to herbaceous and woody plants found on the forest floor. Past research has shown that silvicultural methods can be used to eradicate R. maximum, however it is unclear which of these methods is most efficient and what effects other than stem mortality may occur. In this study, treatments involving prescribed fire, mechanical cutting, and herbicide applications were applied to R. maximum dominated forests in southwestern Virginia to determine what effect seven different silvicultural treatments had on 1) controlling of R. maximum as a forest weed 2) fuel loading inside of a R. maximum thicket, and 3) canopy tree seedling regeneration. Mechanical cutting treatments were successful in reducing R. maximum basal area per acre; however stump sprouting and increased fuel loading occurred. Herbicide applications were successful in controlling only the smallest diameter class of R. maximum stems. Prescribed fire reduced litter layers and caused delayed mortality on R. maximum stems three years following treatment. Hemispherical photographs taken within each plot showed that silvicultural treatments that successfully increased the amount of light entering each plot were influential in seedling establishment three years following treatments. Results from this study can be used to further perfect silvicultural applications that alleviate R. maximum cover on the forest landscape. / Master of Science

Rosebay rhododendron (Rhododendron maximum L.) is an ericaceous shrub commonly found in riparian areas of the Appalachian Mountains. After more than a century of fire exclusion in the U.S., the distribution of R. maximum and its dominance of forest understories have increased. Rhododendron expansion has caused a decline in overstory regeneration and the potential for dangerous fuel conditions around suburban structures near the wildland-urban interface. The purpose of this study was to determine the effects of seven silvicultural treatments on both the fuel loading within an R. maximum thicket and the control of R. maximum as a forest weed. The final objective of the project was to determine the cost effectiveness of each implemented treatment. Due primarily to moisture conditions, a single prescribed burn was relatively ineffective in reducing fuel loading and causing R. maximum mortality. Mechanical cutting caused a drastic shift in the size-class distribution of R. maximum but resulted in heavy sprouting and increased fuel loading. Herbicide application did not reduce or increase fuel loading and was important in R. maximum control only when combined with other treatments. The prescribed burning treatment was the least expensive individual treatment while mechanical cutting was the most expensive. Combination treatments showed increased effectiveness in controlling R. maximum but were more expensive than the individual treatments. The results of the treatments from this study will be used over the long term to demonstrate to land managers the effects of vegetation control on rhododendron. / Master of Science