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Ecological theory predicts that high environmental heterogeneity causes high biodiversity. Theory further predicts that more biodiversity results in greater ecosystem functioning. These theoretical predictions were evaluated in three studies using grassland restorations from agriculture.

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Tropical soils are a globally important store of terrestrial carbon (C) and source of atmospheric carbon dioxide (CO2), regulated by the activity of soil microorganisms, through the mineralisation of plant residues and soil organic matter (SOM). Climatic warming will influence microbial activity, and this may accelerate the rate of C release from soils as CO2, contributing to alterations in current atmospheric composition, and generating feedbacks to climate change. Yet the magnitude of C loss from tropical soils remains uncertain, partly because we do not fully understand how non-climatic factors – including the chemistry of plant inputs, the availability of soil nutrients and the composition of the decomposer community – will interact to determine the response to changes in temperature. This thesis examines how these factors together regulate the rate of C cycling in contrasting soils across a 3400 m tropical elevation gradient in the Peruvian Andes, spanning a 20 ºC range (6.5 – 26.4 ºC) in mean annual temperature. Large-scale field-based manipulation experiments, translocating leaves and soil-cores across the elevation gradient (to impose an in-situ experimental warming treatment), were combined with controlled laboratory studies to examine the microbial-scale mechanisms which underlie the processes of decomposition and soil respiration observed in-situ. Results show that, across the gradient, rates of leaf-decomposition were determined principally by temperature and foliar chemical traits, while soil fertility had no significant influence. The effect of temperature was, however, stronger across higher-elevation sites, suggesting a greater vulnerability of the C-rich soils in montane systems to increased C loss under climatic warming. In lowland forests, the presence of invertebrate macrofauna also accelerated rates of decomposition, but leaf chemistry explained the greatest proportion of the observed variance, with a strong role for leaf chemical traits also identified under controlled conditions. Despite marked differences in microbial abundance and community composition among soils, these metrics were not associated with observed rates of decomposition. These results suggest that climate-related changes to plant species distributions (with associated changes to the chemistry of leaf-inputs), and upslope extension of macrofaunal ranges, could strongly influence future rates of leaf decomposition, independently of the direct response to warming. From the soil translocation study, root-soil interactions stimulated substantial net C loss from montane soils following translocation downslope (experimental warming treatment), indicating that warming-related changes to root productivity, exudation and/or species-composition could represent an important mode of future C loss from these soils. To examine more closely how inputs of plant-derived C influence the turnover of pre-existing SOM, and whether soil nutrient availability modulates the response, soils were amended with simple and complex 13C-labelled substrates in combination with inorganic nutrient treatments. Isotopic partitioning was used to determine the degree to which C and nutrient inputs accelerated (positive priming) or retarded (negative priming) the decomposition of SOM. Amendment of upper montane forest and montane grassland soils with nitrogen (N; alone and in combination with C) substantially retarded the decomposition of SOM, suggesting that microbial demand for N strongly regulates the turnover of organic matter in these soils. In contrast, amendment of lower montane and lowland forest soils with C stimulated positive priming of SOM, which was strongest in response to the simple C substrate and was not influenced by nutrient treatments, suggesting that microorganisms in these soils are primarily constrained by availability of labile C. Functional differences among microbial groups were also evident, with gram-negative bacteria and fungi using more labile sources of C while gram-positive bacteria used more complex C. Together, results from these studies considerably advance our understanding of soil C dynamics across lowland and montane systems, painting a rich picture of interacting processes which will determine the future soil C balance in tropical ecosystems. They show that the influence of temperature on the rate of soil C cycling is strongly affected by the nature and composition of plant-derived and atmospheric inputs, the principal additional constraints varying with elevation, leading to both opposing and reinforcing effects on rates of decomposition. The greater observed temperature sensitivity of decomposition at higher elevations is coupled with high microbial demand for N which regulates the turnover of SOM, whereas at lower elevations leaf decomposition is accelerated by active macrofaunal breakdown, while microbial decomposition of SOM is constrained by the availability of labile C. Under a global change scenario of increased temperature and N deposition, results therefore suggest that: (i) modified chemistry of plant inputs will influence rates of decomposition, independently of climate; (ii) increased availability of labile C will lead to more rapid decomposition of SOM at lower elevations; (iii) greater root productivity (associated with warming and plant-community shifts) will stimulate soil C loss across montane regions; but (iv) at higher elevations, a possible countervailing effect may be imposed on rapid warming-accelerated decomposition if increased N availability reduces microbial mineralisation of SOM. The net effect on the ecosystem C budget will depend on the balance of C gain from primary productivity and C loss from soils. Overall, however, the results presented here suggest that the large soil C stores in higher-elevation montane regions are particularly vulnerable to substantial reductions under exposure to short- and medium-term climatic warming.

Reindeer appear to have strong positive effects on plant productivity and nutrient cycling in strongly nutrient-limited ecosystems. While the direct effects of grazing on vegetation composition have been intensively studied, much less is known about the indirect effect of grazing on plant-soil interactions. This thesis investigated the indirect effects of ungulate grazing on arctic plant communities via soil nutrient availability and plant nutrient uptake. At high density, the deposition of dung alone increased plant productivity both in nutrient rich and nutrient poor tundra habitats without causing major changes in soil possesses. Plant community responses to dung addition was slow, with a delay of at least some years. By contrast, a 15N-urea tracer study revealed that nutrients from reindeer urine could be rapidly incorporated into arctic plant tissues. Soil and microbial N pools only sequestered small proportions of the tracer. This thesis therefore suggests a strong effect of dung and urine on plant productivity by directly providing nutrient-rich resources, rather than by stimulating soil microbial activities, N mineralization and ultimately increasing soil nutrient availability. Further, defoliation alone did not induce compensatory growth, but resulted in plants with higher nutrient contents. This grazing-induced increase in plant quality could drive the high N cycling in arctic secondary grasslands by providing litter of a better quality to the belowground system and thus increase organic matter decomposition and enhance soil nutrient availability. Finally, a 15N natural abundance study revealed that intense reindeer grazing influences how plants are taking up their nutrients and thus decreased plant N partitioning among coexisting plant species. Taken together these results demonstrate the central role of dung and urine and grazing-induced changes in plant quality for plant productivity. Soil nutrient concentrations alone do not reveal nutrient availability for plants since reindeer have a strong influence on how plants are taking up their nutrients. This thesis highlights that both direct and indirect effects of reindeer grazing are strong determinants of tundra ecosystem functioning. Therefore, their complex influence on the aboveground and belowground linkages should be integrated in future work on tundra ecosystem N dynamic.

Abstract Plant community composition and diversity are determined by the balance between rates of immigration and extinction. Processes of immigration to a local community, i.e. propagule availability and dispersal of propagules between and within habitats, set the upper limit for the pool of species potentially capable of coexisting in a community, while local biotic interactions, i.e., competition, facilitation, herbivory and interactions with below-ground ecosystem components, and environmental factors control colonisation and establishment, and determine the persistence and dynamics of already existing species. In this thesis, I studied (1) the interactions between propagule availability, biotic and environmental constraints on colonisation, and (2) the interdependence between biotic and environmental factors regulating community processes in already established resident vegetation. First, I found that both propagule availability and competition with adult plants limited the rates of colonisation and total community diversity in a relatively low-productive tundra ecosystem. Long-term exclusion of mammalian herbivores and alleviation of nutrient limitation by fertilization increased the intensity of competition with established vegetation, and diminished immigration rates. In addition, I also found that community openness to colonization depended on the initial community properties, i.e., the functional composition and the traits of dominant plants in resident vegetation, which mediate the effects of nutrient addition and biomass removal on immigration rates. Second, adult plants in the resident vegetation experienced an increased extent of neighbourhood competition and herbivory in nutrient enriched conditions and in naturally more fertile habitats. However, the effects were also species-specific. On a community level, release from heavy grazing favoured lichens over graminoids and increased species richness. Furthermore, I also showed that plant community composition was strongly linked with soil organic matter quality and microbial community composition, and that these vegetation-soil-microbe interactions varied along a gradient of soil pH. Overall, my results emphasise that propagule availability, biotic and environmental control over community processes are strongly interconnected in tundra ecosystems. Especially, my findings highlight the role of plant competition and herbivory and their dependence on soil nutrient availability in governing colonisation and resident community dynamics. My results also indicate that plant functional composition and traits of dominant plants are of great importance in channelling community responses to external alterations and dictating plant-soil interactions.

As ecosystem engineers, invasive earthworms are one of the main drivers of plant community changes in North American forests previously devoid of earthworms. One explanation for these community changes is the effects of earthworms on the reproduction, recruitment, and development of plant species. However, few studies have investigated functional trait responses of native plants to earthworm invasion to explain the mechanisms underlying community changes. In a mesocosm (Ecotron) experiment, we set up a plant community composed of two herb and two grass species commonly found in northern North American forests under two earthworm treatments (presence vs. absence). We measured earthworm effects on above- and belowground plant biomass and functional traits after 3 months of experiment. Our results showed that earthworm presence did not significantly affect plant community biomass and cover. Furthermore, only four out of the fifteen above- and belowground traits measured were affected by earthworm presence. While some traits, such as the production of ramets, the carbon and nitrogen content of leaves, responded similarly between and within functional groups in the presence or absence of earthworms, we observed opposite responses for other traits, such as height, specific leaf area, and root length within some functional groups in the presence of earthworms. Plant trait responses were thus species-specific, although the two grass species showed a more pronounced response to earthworm presence with changes in their leaf traits than herb species. Overall, earthworms affected some functional traits related to resource uptake abilities of plants and thus could change plant competition outcomes over time, which could be an explanation of plant community changes observed in invaded ecosystems.

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.