Reindeer can have strong effects on the plant species composition and functioning of tundra ecosystems, and often promote a transition towards a graminoid-dominated vegetation type. As a result, they influence many ecological processes, such as nutrient dynamics, soil biotic composition and functioning, and carbon storage. Several studies suggest that the effect of reindeer on vegetation may follow predictable patterns and could induce an alternative stable vegetation state. However, little empirical data on the long-term stability of reindeer effects on vegetation exist, as it is inherently challenging to study these ecological processes experimentally on a sufficiently long timescale. The main objective of this thesis was therefore to gain a better understanding of the long-term ecological processes following reindeer-induced vegetation shifts. In order to gain a more mechanistic insight in what initially drives this transition, I used a field-based grazing simulation experiment in which I separated defoliation, trampling, moss removal and the addition of feces. This allowed me to test the relative contribution of reindeer-related activities to initiating the shift from moss and heath- dominated tundra towards a graminoid-dominated vegetation state. Additionally, I studied the long-term ecological stability following such a vegetation shift. I did this by addressing historical milking grounds (HMGs): sites where high reindeer concentrations associated with historical traditional reindeer herding practices induced a vegetation transition from shrubs towards graminoids several centuries earlier, but which were abandoned a century ago. Studying HMGs allowed me to address: 1. The potential stability of reindeer-induced vegetation shifts; 2. The ecological mechanisms contributing to the long-term stability of these vegetation shifts; and 3. How such long-lasting vegetation changes influence soil carbon- and nutrient cycling. I found that trampling by reindeer is an important mechanism by which reindeer cause vegetation change. Addressing HMGs further revealed that this vegetation change can be hightly persistent, as the studied HMGs showed only a low encroachment at the surrounding borders in the last 50 years. The vegetation in the core areas of all studied HMGs had remained strikingly stable, and were hardly invaded by surrounding shrubs. Interestingly, soil nutrient concentrations and microbial activities were still different from the surrounding area as well, and even comparable to actively grazed areas. Even after many centuries of changed vegetation composition and soil processes, there was no difference in total carbon sequestration. This suggests that the environmental conditions for microbial decomposition were more important than vegetation composition for the soil carbon stocks, in our study site. After studying the contemporary habitat use of HMGs by reindeer and other herbivores, investigating the potential plant-soil feedbacks mechanisms and detailed soil analyses, I concluded that several ecological mechanisms contribute to the long-term stability of HMGs: first, the altered soil biotic and abiotic conditions appear to have a stronger advantage for HMG vegetation than for the surrounding tundra vegetation. Furthermore, I found a clear browsing preference of small rodents on single shrubs proliferating in HMGs, causing a strong limitation on shrub expansion. Moreover, the dense established sward of graminoids likely poses a strong direct competition for space and nutrients, hindering seedling establishment. Finally, I conclude that HMGs are highly stable on relevant ecological timescales, and propose how the concepts of historical contingency and ASS can be applied to understand stability of these reindeer-induced vegetation transitions.

<p>Eastern black walnut (<i>Juglans nigra</i> L.) ranks among the most highly valued timber species in the central hardwood forest and across the world. This valuable tree fills a critical role in native ecosystems as a mast bearing pioneer on mesic sites. Along with other <i>Juglans</i> spp. (Juglandaceae), <i>J. nigra</i> is threatened by thousand cankers disease (TCD), an insect-vectored disease first described in 2009. TCD is caused by the bark beetle <i>Pityophthorus juglandis</i> Blackman (Corthylini) and the phytopathogenic fungus <i>Geosmithia morbida</i> Kol. Free. Ut. & Tiss. (Bionectriaceae). Together, the <i>P. juglandis</i>-<i>G. morbida</i> complex has expanded from its historical range in southwest North America throughout the western United States (U.S.) and Europe. This range expansion has led to widespread mortality among naïve hosts <i>J. nigra</i> and <i>J. regia</i> planted outside their native distributions.</p> <p> The severity of TCD was previously observed to be highest in urban and plantation environments and outside of the host native range. Therefore, the objective of this work was to provide information on biotic and abiotic environmental factors that influence the severity and impact of TCD across the native and non-native range of <i>J. nigra</i> and across different climatic and management regimes. This knowledge would enable a better assessment of the risk posed by TCD and a basis for developing management activities that impart resilience to natural systems. Through a series of greenhouse-, laboratory- and field-based experiments, environmental factors that affect the pathogenicity and/or survival of <i>G. morbida</i> in <i>J. nigra</i> were identified, with a focus on the microbiome, climate, and opportunistic pathogens. A number of potentially important interactions among host, vector, pathogen and the rest of the holobiont of TCD were characterized. The <i>holobiont</i> is defined as the whole multitrophic community of organisms—including <i>J. nigra</i>, microinvertebrates, fungi and bacteria—that interact with one another and with the host.</p> <p>Our findings indicate that interactions among host, vector, pathogen, secondary pathogens, novel microbial communities, and novel abiotic environments modulate the severity of TCD in native, non-native, and managed and unmanaged contexts. Prevailing climatic conditions favor reproduction and spread of <i>G. morbida</i> in the western United States due to the effect of wood moisture content on fungal competition. The microbiome of soils, roots, and stems of trees and seedlings grown outside the host native range harbor distinct, lower-diversity communities of bacteria and fungi compared to the native range, including different communities of beneficial or pathogenic functional groups of fungi. The pathogen <i>G. morbida</i> was also associated with a distinct community of microbes in stems compared to <i>G. morbida</i>-negative trees. The soil microbiome from intensively-managed plantations facilitated positive feedback between <i>G. morbida</i> and a disease-promomting endophytic <i>Fusarium solani</i> species complex sp. in roots of <i>J. nigra</i> seedlings. Finally, the nematode species <i>Bursaphelenchus juglandis</i> associated with <i>P. juglandis</i> synergizes with <i>G. morbida</i> to cause foliar symptoms in seedlings in a shadehouse; conversely, experiments and observations indicated that the nematode species <i>Panagrolaimus</i> sp. and cf. <i>Ektaphelenchus</i> sp. could suppress WTB populations and/or TCD outbreaks.</p> <p>In conclusion, the composition, function, and interactions within the <i>P. juglandis</i> and <i>J. nigra</i> holobiont play important roles in the TCD pathosystem. Managers and conservationists should be aware that novel associations outside the host native range, or in monocultures, intensive nursery production, and urban and low-humidity environments may favor progression of the disease through the effects of associated phytobiomes, nematodes, and climatic conditions on disease etiology. Trees in higher diversity, less intensively managed growing environments within their native range may be more resilient to disease. Moreover, expatriated, susceptible host species (<i>i.e.</i>, <i>J. nigra</i>) growing in environments that are favorable to novel pests or pest complexes (<i>i.e.</i>, the western U.S.) may provide connectivity between emergent forest health threats (<i>i.e.</i>, TCD) and native host populations (<i>i.e.</i>, <i>J. nigra</i> in its native range).</p>