Within the field of ecosystem science, substantial progress has been made towards our knowledge of the factors which shape the global distribution of vegetation. However, factors which control the biogeography of belowground vegetation structure and function remain less understood than their aboveground counterpart. Vegetation types can differ substantially in terms of belowground processes such as root growth, root turnover, and resulting vertical root distributions. Fine roots provide an exchange surface, allowing transport of water and nutrients to the leaves. On the other hand they also represent a significant sink for photosynthetically fixed carbon to the soil in terms of maintenance and growth. Overall, root processes have a major influence on fluxes of water, carbon and nutrients within ecosystems. In this thesis, an electrical impedance method was used to determine the area of ‘active’ root in contact with the soil for the purpose of absorption. These measurements were compared to the leaf area of the trees, for the first time allowing the aboveground and the belowground resource exchange areas of plant to be contrasted. This approach was first developed to compare the exchange surface areas of leaves and roots within a Sitka spruce (Picea sitchensis) managed forest, making measurements in adjacent stands of differing tree density, but identical in age. Stem density was found to significantly influence the proportion of absorbing root area relative to leaves. Following the successful test of the method, it was used to compare the resource exchange areas of eight stands of forest and savanna vegetation in central Brazil. Across a broad gradient of vegetation structure, the results showed progressively more investment in fine root area relative to leaf area across the transition from dense forest to open savanna. However, a contrasting result showed that the forests had a higher absorbing root area to leaf area ratio than savannas. Furthermore, these measured ratios were strongly correlated with tree height across the eight structurally contrasting stands. It appears that absorbing root area index provides a physiologically meaningful way of characterising belowground water uptake ability, it is possible that excessive investment in fine root area, relative to leaf area, may reflect differences in the requirement for nutrient uptake in poor soils. Complementary to the analysis of root absorbing area, measurements of root activity and belowground carbon cycling were made by focussing on two of the eight tropical study sites. Here, the carbon costs of root growth and respiration were quantified to develop a belowground carbon budget for two structurally contrasting Brazilian savannas, using soil respiration measurements and a root presence/absence manipulation experiment. Annual estimates showed that at least 60% of the total CO2 efflux from the soil was contributed by autotrophic processes, with this value rising to 80% during the dry season. Seasonal fluctuations of soil respiration were strongly correlated with soil moisture for both the autotrophic (R2=0.79, pvalue< 0.05) and heterotrophic (R2=0.90, p-value<0.05) components, with maximum flux rates corresponding with 16.4 and 17.7% soil moisture content respectively. Furthermore, autotrophic respiration was found to varied with phonological patterns of fine root growth (R2=0.80, p-value<0.05). It follows that, the way in which phenological processes respond to a changing climate is of potential importance within seasonally dry regions. Diurnal fluctuations of heterotrophic CO2 efflux were correlated with soil temperature (R2=0.74, p-value<0.05), demonstrating a Q10 value of 1.6 across both sites. In contrast, total soil CO2 efflux was not correlated with temperature (p-value=0.31), suggesting that autotrophic respiration is predominantly limited by substrate supply.

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A crucial question in the debate about reintroducing elephant culling is whether the long-term effects of elephants and mesobrowsers on woodlands are similar. Sufficiently high biomass-densities of mesobrowsers may, following reduction or removal of elephants, continue to heavily impact earlier life-history stages of a similar suite of woody plants that elephant impacted, preventing these species from maturing. Thus a similar end-point for woodland structure and composition is achieved. No study exists in the literature where woody plant and habitat utilisation of the savanna browser guild has been determined in the same locality over the same period. A review of 49 years of literature implied that the two groups impact the same core woody-species in the same habitats. Dietary and habitat utilisation of guild members was determined in Hluhluwe-iMfolozi Park, South Africa. A small suite (n = 8) of woody species formed the core diet of all guild members. Herbivores’ densities were determined using a novel GIS approach; all members of the guild showed extensive overlap in habitat use. GPS collars and a GIS were then used to detect zones of different density of impala in the landscape, thus defining, for the first time, a natural fine-grain browsing gradient. Densities of woody seedlings were significantly less (average 48% reduction) in areas of high versus low impala density. A simple browse-browser model, incorporating, in a novel approach, functional groups of plant species, was parameterised from these results and an extensive review of the literature. Outputs suggest that over the long-term (100 years), impala will have a similar impact on woodland structure as elephant. An apparently strong synergistic effect between impala and elephant impact, suggests that reduction or removal of either impala or elephant will radically reduce long-term destruction of woodlands. In smaller or medium sized reserves, where control of mesobrowser populations is practical, profitable and more acceptable than elephant culling, these findings imply a re-direction of management efforts. Management should consider the biomass-density of both groups, rather than just focus on the system’s perceived ‘keystone’ species. Such principles may also apply to temperate and other systems.

Abstract Exotic plant species may facilitate their invasion into native communities through the modification of ecosystem disturbances such as fire regimes. Where frequent fires are common, invasive plants that suppress fire may induce a positive feedback which further suppresses fire and promotes their continued invasion. In the pine rockland savanna ecosystem of south Florida, the frequent understory fire regime may be altered by the fire-resistant invasive shrub Brazilian pepper (Schinus terebinthifolius). In this thesis, I document the interaction of Brazilian pepper and fire in these savannas. I show that fire causes significant (30%-50%) mortality among low-density populations of Brazilian pepper. However, Brazilian pepper exhibits rapid growth and reproduces quickly following fire, and in the absence of fire it has a low mortality rate. Furthermore, Brazilian pepper can cause a reduction in fire temperature from 47° C at low densities, to almost 200° C at high densities, where it can completely impede fire spread. This creates the potential for Brazilian pepper to initiate a fire-suppressing feedback if it can reach a density threshold during extended fire-free intervals. At a landscape scale, I analyzed digital aerial photographs to show that fire frequency correlates with the extent of Brazilian pepper invasion into pine savanna fragments in southern Florida. In savannas where fire is frequent, Brazilian pepper does not heavily invade, but savannas that are heavily invaded tend to be unburned for more than 20 years. This supports both the regulation of low-density Brazilian pepper populations by fire, and the potential for high-density Brazilian pepper populations to suppress fire and facilitate further invasion.

D.Phil. / What underlies species distribution and species coexistence has long been of key interest in community ecology. Several methods and theories have been used to address this question. However, it still remains a controversial debate. The recent development of plant DNA barcodes with possibility of merging phylogeny with ecology brings high expectation in uncovering the processes underlying community assemblages. Previous works that used molecular approach in community ecology focused mainly on rainforests. Using a phylogenetic approach, this study brings novel understandings about savanna ecology, especially regarding how megaherbivores impact plant community composition. The Kruger National Park (KNP) is one of the world’s largest reserves, but less studied from a phylogenetic perspective. A DNA database of 445 DNA sequences (plant DNA barcodes, rbcLa + matK) was generated for the woody plants of the KNP. This database proves reliable in reconstructing the phylogeny of Angiosperms of the park. Based on this phylogeny, the present study characterised plant community composition, and investigated how megaherbivores influence this composition. Results indicate that plant communities in the KNP are not neutral, i.e. they are more clustered than expected under various null models. This suggests that ecological forces, most likely habitat filtering may be playing key role in dictating community structure in the KNP. The KNP is well-known for its richness in megaherbivores. The contribution of these animals to the current shape of plant community structures was therefore further investigated. Where megaherbivores have been excluded, plant diversity decreases, but shifts in plant community structure are contingent upon the initial community composition, suggesting that herbivory might be important filter that drives the clustering pattern observed.

The objective of this study is to determine whether changes in arthropod community structure in restored longleaf pine savannas corresponds to differences in vegetation structure often associated with burn frequency. Longleaf pine savannas are fire-maintained ecosystems characteristic of the southeastern United States and have experienced severe declines (around 97%) since European settlement. Changes in fire regime have been instrumental in the declines. Restoration of these ecosystems has involved reinstitution of periodic burnings to promote and maintain vegetative characteristics of the savannas. This study investigates trends in arthropod communities from areas heavily invaded by hardwood shrubs against those dominated by longleaf pines and associated vegetation. These data suggest that herb-dominated sites have higher overall diversity. While overall abundance differences were not found, significant differences have been detected at the order and family level, indicating that vegetation structure and periodic burning are important factors in maintaining arthropod communities characteristic of these savannas.

M.Sc. / Boreholes in the Kruger National Park (KNP) was at first developed to increase the number of animals that were at low densities because of poaching, diseases, fencing and low permanent water availability. This development of artificial waterholes without managerial measures or knowledge of possible consequence led to negative impacts on the environment. These included that the higher concentration of herbivores around artificial waterholes led to a change in vegetation; secondary vegetation growth did not support the feeding habits of water dependent species; the infiltration rate of water in the soil surrounding waterholes changed and animal distribution patterns also changed, in turn changing the predatory base. Due to the above the KNP developed a new water policy. The policy states that all artificial waterholes that are open should be part of natural ecosystem principles. This led to the closure of many artificial waterholes that did not conform to the requirements of the new water policy. This study aimed to determine the ecological implications of artificial waterholes and whether tourism will be affected by the closure of these artificial waterholes. The following two hypothesis were thus tested in this project: i) Artificial waterholes have ecological implications on the environment. ii) The closure of artificial waterholes will have a negative response from tourists and thus affect tourism to the KNP. The results from this project indicated that both these hypothesises can be accepted as the available literature clearly showed that artificial waterholes do have a negative impact on the environment and that the majority of the tourist questioned asked for the waterholes to be opened again. The latter reaction was mainly due to the tourist‟s concern that the animals will suffer and die without water. Furthermore the visitors are concerned they won‟t see animals anymore. This result can partly be due to the low awareness of the visitors on the subject. No effort was made to communicate these decisions, and the reasons it‟s based on, to the public. Although a certain number of artificial waterholes were closed according to the new KNP policy, it is still important to keep a number open due to fact that KNP is not a natural system. It is suggested however that the closure of the artificial waterholes take place differently, following a principle of rotational opening and closing of patches of waterholes to allow recovery of vegetation and facilitate migration between waterholes.

This study focused on how changes in food availability during the dry season, influenced various aspects of sable (Hippotragus niger niger) foraging and social ecology. The main purpose of this was to determine whether differences in their foraging ecology explained the reduction in population size compared to other more abundant herbivores (e.g. zebra, buffalos, hartebeest). Contrary to expectation, sable did not limit their foraging to woodlands, but also fed in open grasslands, provided these retained green grass during the dry season. Wetlands were key resource areas during the dry season, but sable did not limit their feeding to these bottomland areas. They also fed on hill slopes and upper plateaus that had been burnt. During the study, sable were attracted to burnt areas despite the reduced grass availability. My study highlights how these burnt areas were important in alleviating nutritional deficiencies during the dry season. At lower spatial levels, I looked at the factors that influenced the selection of feeding areas and plant species. Like other grazers, sable were attracted to green leaves but did not avoid brown leaves or stems. Grass species eaten were the same as those preferred by most domestic and wild grazers. Surprisingly, adult males stayed with the breeding herds despite their different activity budgets. Unlike most other African grazers, sub adult sable males did not form bachelor groups. Instead, they stayed within the breeding herds the whole time and had a similar activity budget to females. As adult males moved with the breeding herds, I was able to examine the costs and benefits involved with such a social structure. From this, I was able to relate my results to the bigger picture of factors influencing sexual segregation in ungulates. In conclusion, sable utilised a wide variety of habitat types and adjusted their foraging behaviour to accommodate different seasonal situations. I was unable to find anything in their feeding ecology that could explain the continued decline in population size. Therefore, I suggest that future research needs to focus on other potential causes such as a higher susceptibility to predation and disease as compared to more abundant herbivore species.

Orientador: Davi Rodrigo Rossatto / Resumo: O Cerrado pode estar exposto a diversos distúrbios naturais moduladores, entre eles a geada. Este é um fenômeno capaz de afetar a distribuição e seleção de características de tolerância e sobrevivência das espécies nestas vegetações. Deste modo, o presente estudo pretendeu compreender os efeitos causados pela geada em espécies afetadas e não afetadas por este fenômeno, assim como quantificar características morfo-fisiológicas que possam explicar as respostas de rebrota a baixas temperaturas. As pesquisas foram realizadas em três locais distintos: em Águas de Santa Bárbara - SP na Estação Ecológica de Águas de Santa Bárbara – EecSB (acometida em 2016 por seis episódios de geada entre os meses de junho e agosto) (Capítulo I); Assis - SP na Estação Ecológica de Assis –EEA que foi acometida por eventos de geada no mesmo ano (Capítulo I e II) e na Reserva Ecológica do IBGE em Brasília – DF (Capítulo III). No segundo capítulo as coletas e experimentos foram realizados na EEA e no câmpus da Unesp. No capítulo I e II foram avaliados indivíduos de espécies previamente selecionadas após a ocorrência da geada, com o intuito de analisar os efeitos da mesma na mortalidade de indivíduos e ramos de espécies arbóreas consideradas como afetadas pelo fenômeno; também foi verificado se o espécime teve rebrote a partir do solo ou do caule/ramos e o grau de proteção das gemas, além disso, realizamos análises de respostas morfo-fisiológicas por meio da avaliação de atributos foliares, como, espess... (Resumo completo, clicar acesso eletrônico abaixo) / Abstract: The Cerrado can be exposed to several natural disturbances, being frost one of them. Frost is a phenomenon that can affect the distribution and selects tolerance and survival characteristics of species occurring in these vegetations. This study aimed to understand the effects caused by frost on the biomass of frost affected and non-affected tree species, and to quantify morpho-physiological characteristics that may explain the regrowth responses after the exposition to low temperatures. Data collection was carried out in three different locations: the first chapter was developed in Águas de Santa Bárbara- SP at the Ecological Station of Águas de Santa Bárbara – EecSB, which was affected in 2016 by six episodes of frost between the months of June and August; data for first and second chapter were collected in Assis – SP at the Ecological Station of Assis –EEA, which was also affected by frost events in the same year. The chapter III was conducted at the IBGE Ecological Reserve in Brasília – DF. In the chapter II the collections and experiments were carried out in Assis in the EEA and in Unesp campus. In chapter I and II, individuals from preselected species were evaluated to analyze the effects of frost on the branch mortality of tree species considered to be affected by the phenomenon; additionally it was verified whether the specimen had regrowth from the soil or the stem / branches and the degree of protection of the buds. We performed analyzes of morpho-physiological respo... (Complete abstract click electronic access below) / Mestre

A thesis submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfilment of the requirements for the degree of Doctor of Philosophy The support of the Global Change and Sustainability Research Institute (GCSRI) towards this research is hereby acknowledged. Opinions expressed and conclusions arrived at, are those of the author and are not necessarily to be attributed to the GCSRI. February 2017 in Johannesburg, South Africa / Global change, specifically land cover change and climate change, are recognised as the leading drivers of biodiversity loss worldwide. Habitat loss has resulted in a loss of biodiversity and led to significant declines in species populations. Climate change is altering species distributions, ecosystem composition and phenology. Conservation planning is required to offset these dynamic threats to species persistence into the future. Plants form the basis of trophic structure and functioning and may not be able to track changing environmental conditions as well as mobile species. They thus represent an essential starting point for understanding climate change and habitat loss impacts. The patterns and processes which generate and maintain floristic diversity must be explored before global change impacts on these communities can be assessed and planned for at a landscape scale. This thesis investigates the environmental variables structuring indigenous plant community composition, pattern and turnover in grassland and savanna systems in KwaZulu-Natal. The threats posed by land cover change and climate change are explored and a coarse-grained landscape connectivity map developed to impart maximum resilience in order to maintain floristic diversity in the era of anthropogenically induced global change. The environmental variables correlated to floristic pattern and turnover were temperature, soil fertility and precipitation variables. The orientation of the temperature gradient conflicts with the soil fertility gradient, hence species with particular soil requirements will be hampered in their efforts to track the temperature gradient. The gradients were non-linear with turnover highest on dystrophic soils in warm and drier summer regions. The major drivers of land cover change were cropped agriculture, timber plantations (agroforestry), rural and urban development, dams and mines. The drivers of change differed according to land tenure type. The average rate of habitat loss in the province over an 18 year period was 1.2% per annum, levels which are considered unsustainable. A target level of 50% of natural habitat remaining is recommended. Environmental domains were identified using the environmental correlates of plant community composition. These were used to investigate climate change impacts using a collection of downscaled climate models. Conditions suiting savanna species are set to increase at the expense of conditions suiting grassland species raising significant challenges for the conservation of grasslands. Indices of habitat intactness and climatic stability were used to develop a vulnerability framework to guide conservation actions to mitigate global change impacts on floristic diversity. Building on the insights gained from the study, a connectivity map linking protected areas was developed, that if implemented, will maximise the opportunity to maintain floristic diversity into the future. The spatial location of the corridors was prioritised based on broad scale climatic refugia, high turnover areas and important plant areas for endemic and threatened species. The corridors were aligned along the major climatic gradients driving floristic pattern. The corridors represent the most natural and cost-effective way for species to adapt to climate change and persist in the landscape. This thesis provides new insights into two global threats facing plant communities in KwaZulu-Natal and provides a suite of products that inform dynamic conservation planning and directs appropriate conservation action. The results may be used to inform policy and legislation. / MT2017