The state of Florida has one of the most severe exotic species invasion problems in the United States, but little is known about their influence on soil biogeochemistry. My dissertation research includes a cross-continental field study in Australia, Florida, and greenhouse and growth chamber experiments, focused on the soil-plant interactions of one of the most problematic weeds introduced in south Florida, Lygodium microphyllum (Old World climbing fern). Analysis of field samples from the ferns introduced and their native range indicate that L microphyllum is highly dependent on arbuscular mycorrhizal fungi (AMF) for phosphorus uptake and biomass accumulation. Relationship with AMF is stronger in relatively dry conditions, which are commonly found in some Florida sites, compared to more common wet sites where the fern is found in its native Australia. In the field, L. microphyllum is found to thrive in a wide range of soil pH, texture, and nutrient conditions, with strongly acidic soils in Australia and slightly acidic soils in Florida. Soils with pH 5.5 - 6.5 provide the most optimal growth conditions for L. microphyllum, and the growth declines significantly at soil pH 8.0, indicating that further reduction could happen in more alkaline soils. Comparison of invaded and uninvaded soil characteristics demonstrates that L. microphyllum can change the belowground soil environment, with more conspicuous impact on nutrient-poor sandy soils, to its own benefit by enhancing the soil nutrient status. Additionally, the nitrogen concentration in the leaves, which has a significant influence in the relative growth rate and photosynthesis, was significantly higher in Florida plants compared to Australian plants. Given that L. microphyllum allocates up to 40% of the total biomass to rhizomes, which aid in rapid regeneration after burning, cutting or chemical spray, hence management techniques targeting the rhizomes look promising. Over all, my results reveal for the first time that soil pH, texture, and AMF are major factors facilitating the invasive success of L. mcirophyllum. Finally, herbicide treatments targeting rhizomes will most likely become the widely used technique to control invasiveness of L. microphyllum in the future. However, a complete understanding of the soil ecosystem is necessary before adding any chemicals to the soil to achieve a successful long-term invasive species management strategy.

ABSTRACT Alliaria petiolata (garlic mustard) is an aggressive non-native and invasive forb that negatively impacts native arbuscular mycorrhizal communities and inhibits or prevents the growth of native plant species. Invasive species mitigation and management strategies that use native revegetation vary in success. This study focuses on which species naturally regenerate in areas where A. petiolata has been mitigated to help inform restoration efforts. Seedling emergence of species within the seed bank of four plot types (uninvaded, invaded, chemically treated, and mechanically treated) were observed two years post restoration efforts to determine which native species are likely to persist to seedlings following management. Species abundance and percent cover of all plant species were recorded to evaluate the presence of species in addition to plant health and physiological differences. Native species abundance was significantly reduced within herbicide treated plots as compared to untreated and invaded plots (p=0.02). Plots treated with mechanical removal had the greatest percent cover of native plants as compared to all other plot types but were dominated by colonizing species which is typical of a disturbed habitat. Mechanical removal also resulted in a greater abundance and stability among functional groups of native species, than those treated with herbicide. Forb species dominated coverage of plots over other functional groups when treated with herbicide. Alliaria petiolata invasion and management methods significantly impacted forb and graminoid species, as they had significantly lower abundance in plots treated with herbicide. The results demonstrate that the method of removal as well as the presence of A. petiolata affects emergence of plant species from the seedbank. The additional disturbance of mechanical removal may alter successional trajectories following invasion. Herbicide treatment resulted in the most similar species abundance as the uninvaded reference plots, which had the lowest seedling emergence and percent cover.

Thesis (PhD)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: Invasive species are among the most important global conservation threats. Their management is one of the key conservation challenges that will have to be addressed in the next few decades. The study of real invasions and their management in natural ecosystems provides an opportunity to gain important information on theoretical and applied aspects of biological invasions. This project focuses on the broader ecological context of invasive ant management in an ecologically sensitive island habitat. The thesis has three main components: 1) assessing the role of the invasive ant Pheidole megacephala in the ecosystem and evaluating its threat to the system, 2) evaluating a low-impact management program for the ant, and 3) using a community-level approach to assess ecosystem response to ant removal. The ant occupied almost 30% of the island‘s total land area and reached extremely high densities in some areas. The ant was associated with exotic hemipteran scale insects through trophobiotic mutualisms that facilitated high ant and hemipteran abundances. The highly destructive scale insect Pulvinaria urbicola was among the hemipterans that benefited from ant attendance. High levels of hemipteran feeding resulted in dieback of functionally important and threatened native Pisonia trees, which represented a significant threat to the forest ecosystem. A management program was initiated in response to this threat, consisting of baiting with selective hydramethylnon-based bait delivered in bait stations, accompanied by detailed pre-and post-baiting monitoring. The method was highly effective at suppressing the ants, whilst preventing bait uptake by non-target organisms. It was also cost-effective and adaptable to ant density in the field, but was only effective over short distances. The method may be applicable to other sensitive environments with similar challenges. After ant control, the ant-scale mutualism was decoupled and the Pu. urbicola population collapsed. There were variable responses in different taxa to the removal of these highly abundant exotic species, the most important of which was the recovery in Pisonia trees. Shoot condition and foliage density improved and there was a decrease in sooty mold. Herbivory on Pisonia increased due to recovery of native canopy herbivores, but the overall impact was far less than that of the exotic hemipterans. Soil surface arthropods, a group that may have been vulnerable to the treatment method, were unaffected by baiting. Instead, they increased significantly after ant removal, confirming the ant‘s impact on other arthropods. Other ant diversity and non-ant arthropod abundance increased post-baiting, including the endemic ant Pheidole flavens farquharensis and some functionally important insects such as the Indian cockroach. Natural enemies that interacted predictably with the mutualists were influenced by management. Predators of hemipterans increased significantly after ant removal and were instrumental in the scale population collapse, whereas parasitoids of hemipterans that benefited from the mutualism declined. Additionally, groups that were unrelated to the mutualism were indirectly influenced by management. The natural enemy assemblage as a whole showed recovery to pre-invasion conditions. The study shows how widely interconnected and influential the ant was in the ecosystem. It highlights the threat of the species in natural systems as well as the complex responses following invasive ant removal. Yet, it also demonstrates the potential to safely and effectively manage the species, thereby raising the opportunity for ecosystem recovery. / AFRIKAANSE OPSOMMING: Indringerspesies is van die belangrikste globale bedreigings vir natuurbewaring. Hulle bestuur is van die grootste bewaringsuitdagings wat in die volgende paar dekades aangespreek moet word. Die studie van werklike invalle en hul bestuur in natuurlike ekosisteme bied 'n geleentheid om belangrike inligting te verkry oor teoretiese en toegepaste aspekte van biologiese indringing. Hierdie projek fokus op die breër ekologiese konteks van uitheemse mier bestuur in 'n ekologies sensitiewe eiland habitat. Die tesis het drie hoofkomponente: 1) die beoordeling van die rol van die indringer mier Pheidole megacephala in die ekosisteem en evaluering van sy bedreiging vir die sisteem, 2) die evaluering van 'n lae-impak bestuursprogram vir die mier, en 3) die gebruik van 'n gemeenskaps-vlak benadering om ekosisteem reaksie op mierverwydering te assesseer. Die mier het byna 30% van die totale landoppervlak van die eiland beslaan en het in party areas baie hoë digthede bereik. Die mier was geassosieer met uitheemse dopluis spesies in mutualismes wat hoë mier en dopluis getalle gefasiliteer het. Die hoogs beskadigende dopluis Pulvinaria urbicola was een van die spesies wat bevoordeel is deur die mutualisme. Hoë vlakke van dopluis voeding het die terugsterwe van funksioneel belangrike, bedreidge inheemse Pisonia bome veroorsaak, wat ʼn groot bedreiging vir die ekosisteem verteenwoordig het. ‗n Bestuursprogram is geïmplimenteer as gevolg van hierdie bedreiging, wat bestaan het uit selektiewe hidrametielnoon-gebaseerde lokaas wat in die veld geplaas is in lokaashouers, vergesel deur intensiewe monitering voor en na lokaasplasing. Die metode was hoogs effektief in die onderdrukking van die miere en het lokaasinname deur nie-teiken organismes verhoed. Dit was ook koste-effektief en aanpasbaar volgens mierdigtheid in die veld, maar was slegs effektief oor kort afstande. Die metode mag van toepassing wees in ander sensitiewe omgewings met soortgelyke uitdagings. Na mierbeheer is die mier-dopluis mutualisme ontkoppel en die Pu. urbicola bevolking het drasties verminder. Daar was verskillende reaksies in verskillende taxa tot die verwydering van die oorvloedryke eksotiese spesies, maar die belangrikste reaksie was die herstel van Pisonia bome. Spruittoestand en blaardigtheid het verbeter en daar was ʼn afname in roetskimmel. Herbivorie op Pisonia het toegeneem as gevolg van ʼn herstel in inheemse herbivore, maar die algehele impak was veel minder as dié van die eksotiese dopluis. Grondoppervlak gelidpotiges, 'n groep wat kwesbaar kon wees vir die behandelingsmetode, was onaangeraak deur die lokaas, maar het beduidend na mierverwydering vermeerder. Mierdiversiteit het vermeerder en die Seychelles endemiese mier Pheidole flavens farquharensis is hervestig. Ander gelidpotiges het ook vermeerder, insluitend funksioneel belangrike spesies soos die Indiese kakkerlak. Natuurlike vyande wat geassosieer was met die mutualiste is beïnvloed deur die mierbestuur. Predatore van dopluis het beduidend toegeneem na mierverwydering en was hoogs betrokke by die vermindering van dopluis, terwyl parasiete van dopluis, wat voordeel getrek het uit die mutualisme, gedaal het. Daarbenewens is groepe wat onverwant was aan die mutualisme indirek beïnvloed deur mierbestuur. Die algehele natuurlike vyand gemeenskap het herstel na pre-indringing toestand. Die studie toon hoe wydverbind en invloedryk die mier was in die ekosisteem. Dit beklemtoon die bedreiging van die spesies in natuurlike stelsels asook die komplekse reaksies wat uitheemse mierverwydering volg. Tog demonstreer dit die potensiaal om die spesies veilig en doeltreffend te bestuur, en sodoende die geleentheid vir ekosisteemherstel te skep.

<p dir="ltr">Non-native biological invasions are one of the leading concerns for global biodiversity. The establishment of invasive species reduces local biodiversity, shifts species composition, changes successional trajectories, and alters ecosystem functions. This thesis examines two aspects of invasive plants: (1) the distribution and the most important climatic and anthropogenic drivers of invasive trees across the eastern United States, and (2) an evaluation of invasive plant removal and herbaceous recovery from a government cost-share program that provides financial support for invasive plant management by private landowners.</p><p><br></p><p dir="ltr">Our first study focused on identifying the distribution of invasive trees, and the factors associated with their distribution. This is essential to predicting spread and planning subsequent management. Using USDA Forest Inventory Analysis (FIA) data and random forest modeling, we examined the distribution, and variables associated with the distribution, of invasive tree species. Invasive trees were found in 10,511 out of 299,387 FIA plots. Invasive species basal area and density (trees per ha; TPH) were highest within the central and southern Appalachian Mountains, Michigan, the Northeast, and the southern Coastal Plain of the United States. A random forest model of invasive species basal area (R²= 0.47, RMSE = 0.47) and density (R²=0.46, RMSE=0.50) vs. environmental variables found that both invasive basal area and density were most strongly associated with human footprint, followed by various climatic variables. An equivalent model of native tree basal (R²=0.53, RMSE=9.25) and TPH (R²=0.47, RMSE=8.64) found that native tree basal area and density were most strongly associated with aridity followed by various climatic variables. As human footprint increased, invasive tree basal area and density increased. These results suggest that the distribution of invasive trees is reliant on human alterations to forests.</p><p><br></p><p dir="ltr">Our second study focused on Environmental Quality Incentives Program (EQIP), a federal cost-share program that has provided $25 billion of financial assistance to farmers and non-industrial private forest owners. Few studies have examined whether this program facilitates the recovery of the herbaceous layer while decreasing the dominance of invasive plant species. We surveyed the herbaceous layer of EQIP-treated and untreated (reference) forests across three physiographic regions of Indiana. Using non-metric multidimensional scaling (NMDS) ordination and linear mixed effects models, we evaluated the species composition, richness, diversity, evenness, floristic quality index, and herbaceous-layer cover of EQIP and reference sites. We also used linear mixed models to evaluate how EQIP site treatment affected the diversity of native plant species. Sites treated with EQIP contracts typically had significantly higher native species richness, Shannon’s diversity, and floristic quality than reference sites. There were significant separations in species composition between EQIP treated and reference forests state-wide and in the southern non-glaciated region of Indiana, although composition overlapped between EQIP and reference forests. Our study suggests that EQIP-funded treatments promote increased species richness and diversity. However, the persistent overlap in species composition we observed may signify biotic homogenization due to a long-shared history of anthropogenic disturbances between EQIP and reference sites. Therefore, active restoration of the herbaceous layer might be needed to allow a full recovery after invasive removal.</p>