Reproductive studies in invasive plants are necessary for an understanding of their potential to establish and spread in foreign environments. Elaeagnus umbellata Thunb. (autumn olive) is an invasive woody shrub that flowers early in the spring and is often noted for its abundant fruit set. This study examined the reproductive biology of E. umbellata in Illinois, where it is highly invasive. Hand-pollination experiments were performed to determine the breeding system of E. umbellata, and floral visitors were collected to determine its pollinators. Experiments showed that E. umbellata is a predominantly outcrossing species with a self-incompatible breeding system. However, individual variation was detected in several reproductive characteristics. Pollen tube analyses revealed that a small percentage of individuals allow successful self-pollen tube growth, and self-fruit set resulting from automatic self-pollination (autogamy) was relatively high in a few plants. Automatic self-pollination is possible because the male and female parts of flowers mature sychronously, but the likelihood of autogamy may vary among individuals due to variability in the spatial separation of male and female parts (herkogamy). Variability in the incompatibility system and the level of herkogamy may impact the outcrossing rates and reproductive success of individuals. The majority of floral visitors to E. umbellata were generalist pollinators. Frequently visiting bees included small and large species such as native Andrena spp., Augochlorella aurata, Bombus spp., Ceratina calcarata, Xylocopa virginica, and the introduced Apis mellifera. Bombylius major (large bee fly) and the moth Mythimna unipuncta (armyworm) were also frequent visitors. Most of the above insect taxa are pollinators of E. umbellata based on analysis of pollen on insect bodies. E. umbellata is likely to achieve its abundant fruit set where these common pollinators and other E. umbellata are present. However, in my study sites, many individuals experienced low fruit set on branches that were open to pollinator visitation, suggesting pollen limitation may be common in some years and at certain sites. The discovery of autogamous individuals demonstrates that some E. umbellata individuals may be able to establish and spread even when mates or pollinators are limiting.

Reclaimed coal-mined lands in the Appalachian region can be successful established and utilized for beef cattle production. Currently, these areas are underutilized partly due to an increase in invasive plant species, such as multiflora rose (Rosa multiflora Thunb. Ex Murr.), autumn olive (Elaeagnus umbellata Thunb.), and sericea lespedeza (Lespedeza cuneata (Dum.-Cours.) G. Don). The steep topography and low economic returns from beef cattle make conventional control methods inhibitive. Goats are effective browsing on invasive plant species. An experiment was conducted in 2006 and 2007 at the Powell River Research and Education Center near Wise, VA (77° 43' 30" west longitude, 38° 57' 30" north latitude, elevation 155.5 m) to determine the effects of an ungrazed control, cattle alone grazing, and mixed grazing goats with cattle on forage biomass, botanical composition, relative plant abundance, and animal performance. The three treatments included an ungrazed control, cattle grazing alone, and mixed grazing goats with cattle. Experimental design was a randomized complete block design with two replicates for the control and three replicates for the grazed treatments. Three times during the grazing season the following were measured, analyzed or assessed: nutritive values of pasture, autumn olive, multiflora rose, and sericea lespedeza were assessed; forage biomass was determined by clipping four 0.25 m² quadrants per control replicate and eight 0.25 m² quadrants per grazed replicate; botanical composition and relative abundance of plant species was assessed by the Double DAFOR method from five fixed points in each control replicate and ten fixed points in each grazed replicate; animals were weighed; autumn olive shrub height was measured with a clinometer from a distance of 10 m from the shrub. Branch length was measured with a tape measure from the base of the branch to the end tip. Shrub survival was measured by counting shrubs in each replicate and determining visually percent leaf-out. Each year, control and cattle alone treatments had greater (P < 0.05). Generally, grass content increased in the grazed treatments from spring to fall while weed content increased in the control treatment (P < 0.05). By the end of the two experimental years, the legume components of the pasture were low. This was more evident in the ungrazed control than the grazed treatments. The relative abundance of tall fescue and orchardgrass (P < 0.05) increased in grazed treatments while sericea lespedeza became a dominant weed in the control (P < 0.05). Goats showed high preference for sericea lespedeza and maintain this plant in a leafy, vegetative stage. This leafy, vegetative growth was found to acceptable to cattle. Cattle performance was not affected by treatment (P < 0.05) but total animal output was higher for mixed grazed compared to cattle alone treatments (P < 0.05). The nutritive values of multiflora rose, autumn olive, and sericea lespedeza were higher than pasture in most instances (P < 0.05). In our experiment, autumn olive was severely impacted by goat browsing. Shrub survival was lower in mixed grazing (61%) by the end of the experiment compared to over 90% for the control and cattle grazing treatments (P < 0.05). Overall, pastures were utilized more uniformly in mixed grazing compared to other treatments. Mixed grazing goats with cattle appear to be a viable option for livestock producers in the Appalachian coal mining region. / Master of Science

AN ABSTRACT OF THE THESIS OF Joshua B. Nickelson, for the Masters of Science degree in Forestry, at Southern Illinois University Carbondale TITLE: EVALUATING THE SUCCESS OF OAK AFFORESTATION ON FORMER AGRICULTURAL FIELDS IN SOUTHERN ILLINOIS Major Professor: Dr. Eric Holzmueller The establishment of oak (Quercus spp.) plantations has increased over the past two to three decades to reduce fragmentation and promote wildlife habitat throughout the Midwestern United States. However, influences such as competing vegetation, previous land cover, plantation size, and site preparation techniques may have varying outcomes on restorative successes. We established 219 plots (.02 ha) in 29 oak plantations located within Crab Orchard National Wildlife Refuge (Williamson County, Illinois) 15-18 years after mechanical planting. Sampling data for all trees over breast height included species, diameter, and lianas existence on the main bole of the tree. Additionally, free-to-grow status was recorded for all oak saplings and estimated cover of the exotic invasive shrub Elaeagnus umbellata and vine Lonicera japonica were documented. Results show significantly higher numbers of total oaks and free-to-grow oaks in plantings previously cropped in clover and soybeans when compared to the fallow sites host to brush species that received treatment (mowing and or herbicidal application). Significantly less oaks in the soybean and clover categories possessed a vine on the main bole of the tree when compared to the treated brush sites. Brush sites showed a significantly less number of total trees compared to clover and soybean covers and a higher percent of autumn olive cover (%) compared to soybeans. No significance was found in the percent of oaks with a vine, the percent of oaks overtopped, E. umbellata density or L. japonica cover (%) across the four previous vegetation categories. One treatment of pre-planting mowing and herbicidal application is not effective on fallow sites that are host to early successional species and money should not be invested on Quercus trees or mowing and herbicidal treatments in these scenarios. The results suggest that it is best to plant Quercus species immediately following clover or soybean harvest on abandoned agricultural lands before early successional species become established.

Autumn-olive (Elaeagnus umbellata Thunb.) is an exotic species that was introduced in the U. S. in the 1800's and widely promoted as an ideal plant for erosion control, wildlife habitat and soil remediation. N-fixation by autumn-olive through a symbiotic relationship with actinomycete Frankia, can alter nitrogen cycling and potentially impair water quality through nitrate leaching. Furthermore, legacy effects of accumulated N following vegetative management and restoration efforts are unknown in areas invaded by autumn-olive. The first objective of this research was to determine if there was a relationship between autumn-olive cover and stream nitrate-N (NO3--N) concentration in twelve forested headwater watersheds in southern Illinois. Secondly, changes in autumn-olive cover and stream nitrogen concentrations were assessed by comparing current results to data collected 6 years prior. Nine of the study watersheds had significantly greater autumn-olive percent cover in 2012 compared to 2006 and mean stream NO3--N concentration significantly increased from 2006 to 2012 in all watersheds. Also, a significant exponential relationship was found between stream NO3--N concentration and autumn-olive percent cover. The long term effects of autumn-olive management on N cycling were also investigated by implementing three different vegetation treatments on invaded areas: cutting of autumn-olive trees (CU), cutting and stump herbicide application (CH) and a no treatment application or control (CO). Treated plots showed that soil NO3--N increased temporarily due to the disturbance. However, after the initial post treatment period of one year, soil water NO3--N concentrations on treated plots shifted, so that CH plots had significantly lower levels of NO3--N than in CO plots. Soil water NO3--N in CU plots also decreased, but it was not significantly different from CH plots. Subsequently, soil water NO3--N declined on CH plots, reaching the lowest levels in the third and fourth year after treatment. Reduction in soil water NO3--N in CH plots was not accompanied by differences among treatments in soil N mineralization rates or soil C:N ratios indicating persistent high nitrification rates contributing to mineralization under treated and control plots. These results demonstrate that encroachment of autumn-olive can transform N cycles in natural areas and affect water quality by saturating the soil with mobile forms of N that are leached out of the system. Significant rates of soil N cycling can persist for years after removal of autumn-olive due to the cycling of legacy N in soils and litter. Nevertheless, the cut and stump herbicide application treatment produced positive vegetation management results by halting N-fixation, controlling autumn-olive re-sprouting and by gradually decreasing N availability and NO3--N flushing in treated areas. Treatment of autumn-olive should have an impact on stream water quality in headwater watersheds, given the significant observed relationship between autumn-olive cover and stream nitrogen levels.

The tocopherol (Vitamin E) content of forage from three invasive shrub species was measured to assess the value of the shrubs as a source of vitamin E for goats browsing on overgrown Appalachian pastures. Plant leaf clusters were collected from multiflora rose (Rosa multiflora Thunb.), autumn olive (Elaeagnus umbellata Thunb.), and Morrow's honeysuckle (Lonicera morowii Gray) in replicated plots at a site in southern West Virginia during the 1999 growing season. Alpha-, beta-, gamma-, and delta-tocopherol were extracted with hexane, separated by high performance liquid chromatography on a normal-phase diol column, and quantified. Significant differences (P<0.001) in concentration were found among species for all forms of tocopherol. Alpha-tocopherol predominated, accounting for more than 90% of the total tocopherols in all three species. Alpha-tocopherol levels increased in all species with maturity; however, the magnitude of the increase was not the same in all species. At the end of the growing season, autumn olive had the highest levels of alpha-tocopherol (1270 ± 55 ppm dry matter [DM]), followed by Morrow's honeysuckle (840 ± 55 ppm DM), and multiflora rose (610 ± 55 ppm DM). Goats grazing on mature browse may obtain adequate intake of vitamin E. High nutritive value and/or low concentrations of antiquality factors may not coincide with the high levels of vitamin E found in mature tissue, and the actual vitamin E intake will depend on the feeding behavior of the goat. / Master of Science

Mining has caused ecosystem losses worldwide, with surface mining disturbing >2.4 million hectares in the United States since 1930. The Appalachian region of the US is home to extensive temperate deciduous forests that provide many ecosystem services and economic benefits. However, >400,000 hectares of forest have been lost due to surface coal mining, with most not being restored back to native forests or other productive land uses. These areas are left fragmented, heavily modified, unmanaged, and densely invaded by non-native plants. Autumn olive (Elaeagnus umbellata) is one of the most prevalent invasive species on reclaimed mines in Appalachia and viewed as one of the main hindrances to the successful reclamation of mined land to restore native forests. In order to better assess the impact autumn olive can have on reclamation success, we characterize autumn olive's performance in various reclamation scenarios and also how the management of autumn olive affects hardwood tree establishment. We review how exotic species impact restoration outcomes, and advocate for a better understanding of how these species could contribute towards a more ecological understanding of reclamation. Reclamation goals are currently assessed after 5 years, prioritizing short-term goals (e.g. erosion control) instead of longer-term goals such as the return of ecosystem function. With a better understanding of plant function and ecological processes, we hope to continue to advance successful reclamation on surface mined lands. / Master of Science in Life Sciences