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Includes bibliographical references., Transpiration is inevitable during photosynthesis; however, it may also function to cool leaves, transport nutrients and drive nutrient acquisition via mass-flow. In addition to transpiration, plants water fluxes occur through hydraulic redistribution (HR). I hypothesized that an important function of plant water fluxes is to drive mass-flow nutrient acquisition, with flux rates positively correlated with nutrient limitation but not deficiency. To test whether nutrient availability regulates mass-flow, Phaseolus vulgaris was grown with N placed at one of six distances behind a root-impenetrable mesh whilst control plants intercepted the N-source. In plants forced to acquire N through mass-flow transpiration rates were 2.9-fold higher and P and K accumulation was greater compared to control plants. The contribution of nocturnal transpiration and HR to nutrient acquisition was assessed by supplying Aspalathus linearis (N-fixer) with no fertilizer or Na¹⁵NO₃ and CaP/FePO₄ either above or below-ground with varying rates of below-ground irrigation. ²H₂O was used to trace HR. HR by A. linearis accounted for the bulk of surface soil moisture at dawn and responded positively to surface fertilization. In contrast, plants supplied below-ground fertilizer exhibited both HR and nocturnal transpiration with increased ¹⁵N and P acquisition. Finally, to establish whether clay fraction moderates mass-flow P availability, Triticum aestivum was grown with 0, 1, 5 or 10% (w/w) clay combined with either Ca-P, Fe-P or inositol-P. Transpiration and nutrient accumulation were monitored. Plants acquired P through massflow and diffusion. The acquisition of N and P increased from 0 to 5% clay (w/w) due to enhanced moisture retention, but further additions (10%) reduced P-availability (Inositol-P > Fe-P > Ca-P). Overall, this thesis explored and confirmed the relatively novel idea that nocturnal and diurnal transpiration by plants are not merely the consequence of stomatal opening for CO₂ acquisition. Rather nocturnal and diurnal transpiration are regulated by nutritional requirements and serves as a driving force for nutrient transport to roots. Likewise, hydraulic redistribution serves to draw water from deep and wet soil layers to the upper layers, which serves as a means to enable uptake of nutrients from the rich, but often dry, upper soil. Plants may thus be opportunistic in their water uptake, taking it up when it is available in order to improve the acquisition of nutrients through mass-flow delivery. Plants in low nutrient substrates elevated their water fluxes for mass-flow nutrient acquisition. Consequently, plants growing in mesic climates with low clay soils are likely to display greater dependence on mass-flow nutrient acquisition. This might vary between C₃ and C₄ plants, which differ in WUE. Plants may also increase mass-flow nutrient acquisition during inter-specific competition thus reducing investment in root proliferation for nutrient interception. Plants growing in elevated atmospheric [CO₂] with suppressed transpiration could show limited mass-flow nutrient acquisition.

While many stem and bud galls contain chlorophyll, and have the potential to photosynthesise, these insect-induced growths are generally thought to act as strong carbon sinks, manipulating the normal phloem transport of the host plant in order to serve the demands of the galling herbivore. This study investigated the photosynthetic capacity of bud galls induced by the wasp, Trichilogaster acaciae longifoliae (Pteromalidae) in the invasive Acacia longifolia. The role of this photosynthetic activity was examined in terms of its ability to subsidise carbon budgets, as well as to provide O₂ to the larvae and consume CO₂ in the dense gall tissue, thereby maintaining O₂ and CO₂ concentrations within the range of larval tolerance. Galls were found to contain an overall chlorophyll concentration that was less than half that of subtending phyllodes and a maximum stomata! conductance only 16% that of phyllodes. Gas exchange measurements indicated that while photosynthesis never fully compensated for the respiratory costs of the galls, light-induced carboxylation within galls contributed substantially to the maintenance and growth of galls, especially in the early stages of their development. Very low levels of O₂ were found within the larval chamber and internal tissues of galls, and these levels responded only marginally, if at all, to light, suggesting that the photosynthetic activity of galls does not play a critical role in providing 0 2 to the larvae. The percentage mortality and metabolic response of larvae in reaction to various atmospheres of reduced O₂ and elevated CO₂ indicated that larvae were tolerant of hypoxia and capable of rapidly reducing their respiratory rates to cope with hypercarbia, at least over the short term. Sustained metabolic arrest may, however, have toxic consequences for insects, causing cell damage or even death. The photosynthetic activity of galls substantially reduced internal CO₂ concentrations, thus preventing CO₂ from accumulating within galls over prolonged periods. Hence, the capacity of galls to photosynthesise has significant implications for the survival of the developing larvae by reducing the risk of hypercarbic_toxicity and supplying additional carbohydrates to the gall and its inhabitants, thereby creating a favourable microhabitat in which to live.

Globally, increasing land-use intensity has led to more intensive farming practices at the local scale and the loss of non-crop habitats at the landscape scale which may affect various ecosystem services. Insect pollination by wild pollinators is especially affected, but their relative impact and possible interactions have been relatively unexplored. There is also considerable evidence for the negative impacts of agricultural activities and agrochemical use on ecosystem services delivered by natural vegetation, but these impacts have not been assessed for the indigenous crop Aspalathus linearis (rooibos). The study was performed on 13 sites in Nieuwoudtville and Clanwilliam. I found evidence that rooibos pollination is dependent on flying pollinators and that the natural environment is an important provider of ecosystem services to rooibos farmers.

This thesis tests the hypothesis that parasitism in Thesium represents a specialised foraging strategy for dealing with the nutrient poor soils found in the CFR.

Due to the low P nature of soils within the fynbos biome of the Cape Floristic Region, plants have developed numerous mechanisms which enable them to better acquire phosphorous. A number of species have been reported to have specialised root morphologies (root clusters, mycorrhizae) that enhance P uptake. Plants may also down-regulate the uptake of P by decreasing the expression of genes that encode P transporters. Two Fabaceae species, Podalyria calyptrata and Aspa/athus linearis, were grown in hydroponics for 5 months at a low P supply of 4μM and P-depletion studies were conducted thereafter at 5 levels of external P (4, 10, 20, 50 and lOOμM). Growth rates (biomass accumulation) were also calculated as were root to shoot ratios for both species. A. linearis had a higher uptake rate than P. calyptrata. While the fresh biomass growth rate was similar in both species, P. calyptrata had a higher dry weight root to shoot ratio than A. linearis. The results showed that both species exhibited a lack of response to increasing P concentrations and had similar RGRs. Their uptake rates differed significantly (p

Includes abstract., Includes bibliographical references., The Cape Floristic Region lies in the southwestern Cape, South Africa, and is known for its high species richness and endemism. This Mediterranean-type climate region receives winter rainfall that averages between 250 and 650 mm annually over most of the region but reaches an average of more than 1000 mm in the mountains of the southwest and southern Cape.

Preserving the structure and functioning of ecosystems and the benefits they provide to society depends on our understanding of how biodiversity influences ecosystem function and which processes determine the composition and diversity of biological assemblages. Ecological, evolutionary and biogeographical processes interact to determine the genesis, maintenance and spatio-temporal arrangement of biodiversity. Here I investigate these processes in the Cape fynbos of South Africa by examining the diversification, biogeography and community assembly of a clade of reticulate-sheathed Tetraria species (Cyperaceae:Schoeneae).

Numerous lineages of the Western Cape of South Africa show affinities with the flora of tropical Africa and with Australasia. Recent work suggests that most migrations between the Western Cape and tropical Africa occur in a northward direction, and that connections between the flora of regions in the Southern Hemisphere are maintained by wind-assisted long-distance dispersal. The Fluitantes clade of lsolepis (Cyperaceae: Cypereae) is distributed throughout these areas and provides a useful study group to assess the general validity of published biogeographical trends. Furthermore, the cooccurrence of several closely related species in the Cape floristic region allows geographical and ecological patterns to be used for inference of speciation processes in the clade. Sequence data of the ITS and atpi-H gene regions were collected for 82 specimens; these were used to construct haplotype networks and phylogenies. By using the Tristan da Cunha endemics in the genus, as well as results from higher-level studies, a dated phylogeny for the Fluitantes clade was constructed and allowed for ancestral character state optimization of distributions by maximum likelihood. Ecological data were extracted from geographic information systems map to test for environmental differentiation in the Cape taxa. The Fluitantes were found to have originated in the Cape 7 million years ago. From there, they spread east and northwards onto the mountains of East Africa and to the islands of the Indian Ocean. Multiple dispersal events to Australia were recorded. Incongruence between the plastid and nuclear gene trees indicate hybridization to have taken place in Australasia, with possible subsequent speciation. Although the multivariate analysis found some ecological differentiation between the three Cape species, there was substantial overlap in all variables, and interpretation of habitat differences was difficult. It is suggested that, instead, differentiation may have taken place at the microhabitat scale, as I. rubicunda occupies low-lying sandy depressions, I. striata occurs at higher altitudes floating in water, and I. Judwigii inhabits the edges of streams and wetlands.

Includes abstract., Includes bibliographical references (leaves 59-67)., The Proteaceae are a distinctive component of South Africa’s Fynbos and Australia’s Kwongan vegetation and display a number of adaptations to the soils of these nutrientpoor regions, where availability of phosphate (P) is particularly limited. The Proteaceae are notable for the virtually family-wide ability to form cluster roots.

Plant species with small leaves and sclerophylls have been reported to occur mainly on dry, low-nutrient soils in situations of high insolation. However; a number of physiological functions have been proposed for the two sets of traits. Ferns are well-suited to the study of leaf structure and its impact in different environments, as they show remarkable variation in leaf dissectedness and sclerophylly, and are able to inhabit a variety of marginal habitats. In this study, ecological and leaf trait data were collected for 17 fern species occurring in Skeleton Gorge, Table Mountain, in the Western Cape. Correlations between the traits and regressions of leaf traits on potential environmental determinants was carried out using both species averages and phylogenetically independent contrasts. The habitat and leaf traits were also subjected to a test of evolutionary trait conservatism. Sclerophyllous plants were found to be have thicker leaves, containing less chlorophyll, but sclerophylly was poorly correlated with leaf dissection. Plants occurring in high-light environments tended to be more sclerophyllous and have more dissected leaves, although these environments also were nutrient-poor. Leaf dissection appears to be primarily a means of dissipating heat by convection, rather than evaporative cooling, but it may also improve nutrient acquisition in low-nutrient soils. Sclerophylly in these ferns was not directly associated with nutrients; instead low sclerophylly seems to be favoured in low-light environments, perhaps because of lower metabolic costs or to reduce self-shading.

1. Leaf hairs have been associated with reducing environmental stress by increasing reflectance and the thickness of the boundary layer, however, such effects are more obvious in species with dense trichome layers. The adaptive functions for species with sparse leaf hair cover, such as in the Cape Proteaceae species of the Cape Floristic Region (CFR), remain unknown. 2. We postulated that leaf hairs of the Proteaceae in the CFR facilitate both nutrient absorption and water retention on the leaf surface. Leaf hair distribution is expected to show a pattern relative to the higher levels of atmospheric nutrients with proximity to the coast. 3. We followed a three pronged approach, this involved (i) evaluating the relationship of leaf pubescence with leaf surface water holding capacity, water uptake and nitrogen absorption from wet deposition for 12 Proteaceae species from the CFR, under controlled conditions; (ii) evaluating the relationship between leaf pubescence and plant height of 39 Proteaceae species from the CFR and (iii) comparing visual scores of pubescence with the relative proximity of the plants to the coast (oceanic index/continentality) of 205 Proteaceae species. 4. There was a positive correlation between leaf pubescence (0 to 8 x 10⁶ m⁻²) and water holding capacity for the 12 experimental Proteaceae species, however, water uptake did not vary with hairiness. ¹⁵NO₃⁻, ¹⁵NH₄⁺ and ¹⁵N-glycine solutions were each independently applied as wet deposition to the 12 experimental Proteaceae species. The uptake of glycine showed a positive correlation with hairiness overall and for Protea, whereas nitrate and ammonium uptake were positively correlated only with hairiness of Leucospermum. 5. Average plant height was positively correlated with leaf hair density of the 39 Proteaceae species. The visual score of pubescence was significantly different for each genus and there was no correlation between leaf pubescence and distance from the coast for the 205 field Proteaceae species. 6. Leaf hairs probably play a role in reducing environmental stresses, however, it is also possible that they play a role in water retention and nutrient absorption. Hairs on leaves vary in arrangement and morphological characteristics, thus they accordingly, probably serve different functions.

Includes bibliographical references (leaves 85-91)., Many grassland areas throughout the world may support a woody biomass if fire is suppressed. It is puzzling that fire-tolerant savanna trees do not grow in these grasslands. The Highveld grasslands of South Africa are one such grassland. Hypotheses including fire, human intervention, grass competition and various attributes of soil have been proposed to explain the tree-less nature of the Highveld grasslands, but they have mostly been discounted. In this study it was hypothesised that cool temperatures or low nutrient availability would result in slow growth of saplings in grassland areas that would subsequently not be able to escape frequent fires. Alternatively, frost may exclude trees from grasslands. A seedling transplant experiment of savanna tree species of the Acacia genus, into grassland and savanna areas arranged across an altitudinal gradient, was used to compare growth in these varying climates over one growing season, and the influence of frost on seedlings in the following winter. Soils were collected from grassland and savanna regions to establish if nutrients varied between these areas, and seedling growth was measured in a pot experiment including these different soils. To minimise the effect of other variables, seedlings were watered and grass was excluded. Higher altitude grassland areas were cooler, and the grassland soils that were collected were nutrient-poor, relative to the savanna equivalents, with the exception of one nutrient-poor low altitude soil. Growth was well correlated to both temperature and nutrient availability, and in general there was slower growth in grassland climates and grassland soils compared to in savannas. These seedling growth rates were extrapolated to the growth rates of saplings in natural environments and the time it would take saplings to reach a height above flame height was calculated. This showed that although there were significant differences between growth rates in grassland and savanna soils, the magnitude of these differences was not large enough to prevent saplings from growing into adults in grassland soils. Differences in growth due to temperature variations, however, were large enough to suggest that saplings in grassland climates would grow too slowly to ever reach escape height between frequent fires. Frost caused damage to seedlings and decreased seedling survival at the highest elevation sites, but trees were absent well below the altitudinal limit of frost damage. There has been much discussion about the tree-less nature of the Highveld grasslands, but very little experimental work to back it. Neither temperature nor fire alone can explain the lack of trees. This study provides empirical evidence that slow growth due to low temperatures in combination with frequent fire could exclude savanna trees from the grasslands. The effect of grass competition still needs investigation. The incorporation of fire is useful as it is a fundamental part of C4 ecosystems; this, on top of a base of variable growth due to changes in resource availability. The barely recognised savanna tree-line deserves attention, as savannas continue to invade grasslands in a warming world.

Numerous "Fynbos" species of the Cape Floristic Region (CFR) have particularly fine, narrow leaves. The rates of transpiration and heat loss are partially dependent on boundary layer conductance, which is determined by leaf shape and size, surface modifications and wind speed. We expected fine-leaved species with higher boundary layer conductance to transpire faster than broad-leaved species at low temperatures whereas at higher temperatures we expected transpiration to be limited by stomata! conductance. In contrast, the rate of heat loss may be constrained by thick boundary layers in larger leaves at high temperatures. Leaf gas exchange characteristics at various temperatures were correlated with boundary layer thickness, leaf area and specific leaf area for 14 Proteaceae species using phylogenetically independent contrast species. When the temperatures of individual leaves were altered, while ambient temperature was kept at l 8°C, water loss decreased significantly at both 12°C and 30°C with increased leaf size and thus boundary layer thickness. At 30°C, small leaves with thin boundary layers resulted in leaf temperatures below ambient, while larger leaves with thicker boundary layers had leaf temperatures closer to ambient. However, at 30°C the variation in leaf temperature between the smallest and largest leaves was only 3.4°C. Such a small variation in leaf temperature is unlikely to alter temperature-dependent physiological processes. We conclude that the small boundary layer associated with small leaves enables fine-leaved species to transpire at faster rates when water is plentiful. This may be a particularly important strategy for plants that take up most of their nutrients in the wet winter months from nutrient-poor highly leached soils of the CFR region. We suggest that fine leaves are an adaptation for nutrient uptake during winter, although they may also have the benefit of improved coupling of leaf to ambient temperature during the summer drought period.

Includes bibliographical references (p. 100-107), This study investigated the carbon allocation patterns that enable the spindle form of Acacia karroa found in the HluhluweiMfolozi Park, KwaZulu-Natal, to survive repeated and frequent topkill. The central hypotheses were that resprouting is dependent on root carbohydrate reserves and that very frequent topkill would progressively lower the carbohydrate reserves of a plant and eventually cause mortality.

The adaptive significance of variety in leaf forms remains a mystery for many plant groups. This study aimed, using a combined experimental and comparative approach, to investigate the functional significance of leaf size and shape variation, as well as the patterns of leaf form variation in the genus Jamesbrittenia within the context of phylogenetic history. Leaf sizes 63-measured Jamesbrittenia varied between 0.006cm² (in J. microphylla) and 6.52cm² (in J. megaphylla). Correlations between leaf form and the environment suggest leaf size and shape are primarily adapted to water availability, with only leaf dimension being significantly associated with temperature, while, soil fertility shows no relationship with leaf size. Life history is important, however, as broader leaved annuals and species with shorter-lived leaves are associated with more arid habitats. Results of the experimental trials suggest that the primary function leaf size and shape reduction is to reduce water loss, and not to increase heat shedding. While larger leaves transpire more on a leaf-by-leaf basis, transpiration may be higher in broader leaved species at the whole-plant due to higher total plant transpiration. Thus, it is suggested a reduction in leaf size and dimension in Jamesbrittenia is an adaptation to more arid environments. Alternatively, a change in life history may enable a plant to escape harsh periods and capitalise on favourable times. A small- to intermediate-leaved, perennial ancestor is inferred for Jamesbrittenia, which was associated with arid regions in either the summer or winter rainfall regions of southern Africa. Shifts to an annual life history in Jamesbrittenia are associated with a shift to drier habitats, particularly in the arid winter rainfall region of South Africa.

The phosphorus (P) uptake capacities and tolerance to high P of three Proteaceae species from acid (Leucadenron coniferum, Leucadendron salignum) and alkaline (Protea obtusifolia, Leucadedndron meridianum) soils in the Cape Floristic Region (CFR) were compared. These species were also investigated as potential candidates for growth on nutrient enriched, post-agricultural soils. In parallel, two crop plant species (Lupinus albus, with cluster roots; and L. angustifolius, without cluster roots) were also compared. The cluster-root forming species (L. albus) was more sensitive to higher P levels and showed toxicity in terms of photosynthesis, efficiency of PSII, transpiration and stomatal conductance. Growth, gaseous exchange and efficiency of photosystem II (as a measure of stress) were determined at 1, 10 and 100 μg P g⁻¹ sand for both lupins and Proteaceae. Kinetics of P uptake (in hydroponics) was determined at a range ( 4 to 2000 μM P) of P concentrations. Phosphorus sensitivity was relatively high in Ld. coniferum as evident from reduced growth, net photosynthetic rate, stomata} conductance and transpiration rates. Protea obtusifolia showed the least sensitivity to increased phosphorus concentrations under acidic and native alkaline conditions. The inability to regulate P uptake rates at elevated phosphorus concentrations and excessive P accumulation were major determinants of P sensitivity. The relatively higher P sensitivity of Lupinus albus compared to L. angustifolius is likely related to the efficient P uptake by cluster roots in the former.

The aim of this study was to gain a comprehensive understanding of the patterns of carbon partitioning, storage and remobilization in Acacia karroo during the juvenile life history stage. Tuber total nonstructural carbohydrate (TNC) concentrations and δ¹³C values were determined in plants from two different stages in the juvenile life history of A. karroo. These were one year after a fire when the plant consisted of numerous leafy shoots or coppices (coppicing stage) and three years after a fire when the plant consisted of one pole like stem (gulliver stage). Gullivers were found to have mean TNC pools of 150g and mean TNC concentrations of 33%. Coppices had mean TNC pools of97g and TNC concentrations of 24%. Both total TNC pools and TNC concentrations in gullivers were significantly higher (p < 0.05) than in coppices. Carbon isotopes were used to determine whether growth was based on carbon reserves as heterotrophic growth shows a distinct enrichment in δ¹³C. The water relations of plants can also influence the δ¹³C values of plant growth. However, there were no significant differences in root size and depth between the stages, indicating that all plants had access to similar water sources. Mean δ¹³C values from the stems of plants in the gulliver stage were significantly enriched(> 1‰) in ¹³C compared to both coppicing plants (p < 0.01) and adults (p < 0.05). The negative δ¹³C values in coppice stems suggest that their growth is not based on stored carbon. The enriched δ¹³C values found in the gulliver stems support the hypothesis that carbon reserves are utilized to achieve fast growth rates in an attempt to escape the fire trap. However, the small magnitude of the differences in δ¹³C between the two stages suggests post-burn regrowth is derived from both current photosynthate and stored carbon.