Disturbances, such as herbivory and fire are commonplace in savanna ecosystems. The effects of herbivory and fire on growth and defences of adult trees is a much studied filed in plant ecology. However, there are comparatively few studies that have investigated the effects of herbivory and fire on seedling growth, defences and establishment. African Acacia trees are common and widespread, and are suggested to be keystone species in savanna ecosystems. They have been shown to significantly positively increase soil characteristics, such as soil moisture and infiltration, while also increasing spatial heterogeneity of savannas. These trees are distinguishable by their array of physical and chemical defences. Physical defences are either in the form of spines (physiologically costly to produce, thus considered inducible) or prickles (physiologically cheap to produce, thus considered constitutive), while condensed tannins are the most common form of chemical defences. Adult Acacia trees have been shown on several occasions to be highly resilient to disturbance events, primarily due to their large size. However, the effects of herbivory and fire on Acacia seedlings have been little studied despite their apparent importance for our understanding of African savanna ecosystem functioning. In two separate experiments, this thesis aimed to investigate the individual and combined effects of simulated herbivory and fire on the regrowth, defences, and total non-structural carbohydrates (TNC) of the seedlings of several Acacia species, while also testing the Resource Availability Hypothesis (RAH) (14 Acacia species) and the Expanded Growth-Differentiation Balance Hypothesis (GDBH) (three Acacia species). The RAH (Coley, Bryant & Chapin 1985) and the GDBH (Loomis 1932) both assume that arid-adapted (resource-poor) species are slow growing, with low tissue turnover rates. Contrastingly, humid-adapted (resource-rich) species have fast growth rates and thus a high tissue turnover rate. Therefore, arid-adapted species are predicted to invest more carbohydrate reserves in defence after a disturbance event, in order to defend new photosynthetic material. Conversely, humid-adapted (resource-rich) species are predicted to invest carbohydrate reserves into increased growth after a disturbance event, in order to compensate for tissue loss. The first greenhouse experiment found that, in accordance with the RAH, humid-adapted species displayed elevated growth rates compared to arid-adapted species. Overall, defences significantly increased after herbivory, but significantly decreased after fire. Herbivory was also shown to significantly reduce TNC stores in roots more than fire. We found that arid-adapted species did indeed invest more carbohydrate stores (TNC) into defence, and displaying an increase in spine and prickle abundance, spine and prickle length, and condensed tannin concentration. Humid-adapted species displayed an increase in growth rather than in defence, with the majority of species only increasing one defensive trait. Humid-adapted species also displayed significant trade-offs (negative correlations) between new stem growth and defence traits, while arid-adapted species overall did not display any significant trade-offs between stem growth and increased defences. The majority of arid-adapted species also displayed an allometric effect for spine abundance and length, with an increase in one trait led to an increase in the other. Prickles were found to be inducible, despite the assumption that due to low physiological cost, they are no inducible. Herbivory and fire were shown to not be substitutable in their effects on Acacia seedlings in a controlled experimental setting. The second greenhouse experiment tested the Expanded Growth-Differentiation Balance Hypothesis (GDBH) in the seedlings of A. erioloba, A. karroo, and A. nilotica using five levels of nutrient availability. Overall, spine abundance and spine length displayed a unimodal trend in all three species, with spine abundance and spine length being greatest at a nutrient availability of 800 mg/ ℓ. Spine abundance, spine length and condensed tannins increased significantly after herbivory, but were shown to significantly decrease after fire. We found that with an increase in nutrient availability, the growth of stems and roots, along with the production of TNC in roots and stems of all species significantly increased. Our data provide mixed support for the assumptions and predictions of the expanded GDBH. The regrowth of stems and physical defences were consistent with the GDBH. Chemical defences (i.e. condensed tannins) were however, inconsistent with the predictions of the GDBH. We have shown that Acacia seedlings are highly resilient to disturbance events, while the growth responses of Acacia seedlings are underpinned by TNC stores in roots. Simulated herbivory and fire are often substituted for one another in controlled experiments. However, we have shown that the effects of herbivory and fire have significantly different effects on regrowth patterns of Acacia seedlings. / Thesis (M.Sc.)-University of KwaZulu-Natal, Pietermaritzburg, 2011.

The structure and composition of savanna vegetation is influenced by resource availability and disturbance. Grasses, a major component of savannas, influence this resource availability by competing directly with trees for light, water and soil nutrient resources. The direct causes of bush encroachment are not always apparent, but are commonly ascribed to overgrazing and consequent decreased grass competition. The interaction, both above and belowground, between tree and grass seedlings and the surrounding grass sward is dependant on many factors, such as soil depth, seedling species and sward composition. These factors, as well as the presence or absence of defoliation, in the form of grazing or fire dictate whether the system will remain in a transition state as savanna or move towards a stable woodland state. The major competitive effects experienced by the tree seedlings were dependant on grass species and nutrient level. A. nilotica was affected by aboveground competition while A. karroo was affected by belowground competition. E. capensis caused the greatest decrease in A. karroo plant biomass. Both E. capensis and H. hirta had large competitive effects on the aboveground biomass of A. nilotica, while S. africanus had the greatest effect on belowground biomass. Increasing nutrient availability resulted in an increase in the competitive effect exerted on A. karroo, while little to no change was seen in the competitive effect exerted on A. nilotica. Soil depth constrained plant size in both tree species. The intensity of belowground interactions on tree biomass was unaffected by soil depth, while aboveground competition had a significant effect on shallow soils. Belowground competition was also of greater importance than aboveground competition in dictating tree seedling height. Grass seedlings growing on all three soil depths differed in mean mass, with E. racemosa having the least mass and T. triandra having the greatest. Simulated grazing by cutting the surrounding sward resulted in biomass increases in all three grass species. Changes in savanna composition and structure are thus likely to be influenced by initial species composition and soil depth and soil nutrient composition. While grazing creates niches for grass seedling establishment, heavy grazing has been observed to increase grass seedling mortality. Encroachment is thus more likely to occur on intensively grazed shallow and deep soils than on medium depth soils. This highlights the importance of ensuring the grass sward remains vigorous by resting and monitoring stocking rates to ensure veld is not over-utilized. It is then possible to maintain some form of tree-grass coexistence at a level where available grazing is not compromised. / Thesis (M.Sc.)-University of KwaZulu-Natal, Pietermaritzburg, 2008.

The long-term economic viability and ecological integrity of savanna rangelands is being undermined by increasingly dense woody thickets at the expense of palatable herbaceous cover. This process is known as shrub- or bush-encroachment. Bush encroachment is a subset of a broader ecological riddle underlying the coexistence of woody and herbaceous vegetation that has been the subject of many ecological models. The ecophysiological mechanisms and interactions between trees and grasses on which most assumptions of ecological models are premised have seldom been tested empirically. This document synthesises the results of greenhouse and field-based investigations of the underlying ecological mechanisms and ecophysiological interactions between encroaching Acacia mellifera trees and grasses in a semi-arid environment. In a greenhouse study, I determined the contribution of N2 fixation to the N-budget of Acacia mellifera under conditions of both varying N availability and competition from grass. Tree seedlings had longer shoots and greater total dry mass in the absence of grass. The leaf δ15N values were lower with grass than without grasses. Thus, trees were more reliant on N2 fixation in the presence of grasses. N2 fixation may enable the tree seedlings to survive competition with grass at critical and vulnerable developmental stages of germination and establishment. In a field removal experiment, I monitored the growth rates, water relations and mortalities of shrubs around which neighbouring woody plants were removed (target) and control shrubs over three years. Results showed target trees to have benefitted from removal of neighbours, which was manifested in significantly faster growth rates, less negative predawn water potential and a relatively small degree of canopy die-back. Nonetheless, neighbouring trees appeared to prevent the whole plant mortalities resulting from severe environmental stress. Growing in close proximity with neighbours could therefore yield positive and negative ecophysiological effects. In another greenhouse experiment, I tested the effects of the separation of moisture uptake with depth between tree seedlings and grasses on two common substrate types. I also examined the influence of repeated grass clipping on the persistence of soil moisture. Results indicated a three-tier rooting pattern with a top layer exclusively exploited by grasses, an intermediate layer occupied by both grass and tree roots and deeper layers exclusively tapped by trees. Tree seedling biomass was negatively affected by grass competition although the biomass of grass was enhanced in the presence of tree seedlings on sandy substrates only. The repeated clipping of grass benefitted tree seedlings on rocky substrate more than it did on sandy substrate. The effects of heavy grazing on soil moisture availability to woody shrubs and thus bush encroachment may be contingent on substrate type, being more acute on rocky terrains. Grass competition suppresses tree seedlings but the removal of grass by grazing weakens this suppressive effect, particularly on rocky substrates. The insufficiency of space and soil resources on rocky substrates may necessitate increased investment in root biomass by plants. It is not known why grasses have lower densities on rocky substrates than on sandy substrates, but the obstruction by rocks disadvantages grasses against tree seedlings, leaving grasses vulnerable to grazing pressure. This may allow the woody plants on rocky substrates to benefit more from grass removal than on sandy substrates. Root restriction by rock barriers and, perhaps, sparse soil volume further lead to small tree sizes on the rocky substrate. Small shrubs are less likely to compete intensively for resources and cause density-dependent mortality. Intraspecific competition may maintain shrub sizes within the threshold that can be supported by available resource pools. I conclude from my results that the two-layer hypothesis of niche separation between savanna vegetation is valid although there is an overlap in the grass and tree rooting depth/moisture uptake. An additional factor that affects the success of A. mellifera is the substrate.Trees are more dense on rocky substrates but grow larger on sand. I have further shown that A. mellifera trees fix nitrogen when competing with grasses but do not do so when grasses are absent. A mechanistic model of savanna dynamics will need to integrate water use patterns, substrate and nutrients to make effective predictions about encroachment patterns. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2010.

Extensive drought - and elephant-related dieback of Colophospermum mopane and Acacia tortitis, respectively, offered an opportunity for increasing understanding of the causes of drought-related patch dieback, the factors influencing elephant utilization of woody plants, and the response of woody plants to both aforementioned determinants of savanna structure and function. The dendrochronological analysis of both species was undertaken to estimate potential rates of replacement, following extensive mortality. Areas of discrete dieback were compared with adjacent paired areas of 'healthy' vegetation, which revealed, on average, 87% and 13% loss of basal area by mortality, respectively. 'Live' and 'dead' plots did not differ in soil type, topography or mean slope, but differed in vegetation structure, soil surface condition, and soil chemistry. Although there was evidence of self-thinning, neither inter - nor intra-specific competition explained dieback. 'Dead', by comparison with 'live' plots, had changed from functioning as sinks of sediment and water to sources, were less likely to retain water because of a poor soil surface condition, and were predisposed to drought effects because of a greater proportion of fines, and Na concentration. Dieback resulted from insufficient soil water for survival during a drought owing to the development of a dysfunctional landscape during 50 years of livestock ranching. Spatial heterogeneity within a landscape was suggested to enhance woodland resilience to severe droughts by ensuring the survival of plants in run-on sinks or 'drought refuge' sites. Stem sections were removed from 40 multi-stemmed C. mopane trees and prepared for examination under a dissecting microscope. It was impossible to age C. mopane, owing to a hollow and/or dark heartwood. Nevertheless, the distribution of stem diameters suggested a single recruitment event. Fire scars attributed to the last recorded fire in 1948 could explain the trees' multi-stemmed growth form and indicate that most trees of VLNR were > 50 years of age. Growth rings were identified in 29 A. tortilis trees of unknown age, but were not correlated with annual rainfall records. Growth rates varied between trees; mean ring width ranged from 1.4 to 3.5 mm (overall mean 2.4 ± 0.1 mm). A technique was proposed for predicting growth rate from annual rainfall, using selected data, and several factors potentially influencing ring width in semiarid environs were identified. Permanent ground-based transects were located within riparian (n = 16) and Acacia (n = 5) woodlands to monitor elephant utilization. Elephant had not changed the population structure of the woodlands by 2000, but had reduced stem density from 215.6 stems ha -1 (1996) to 84.4 stems ha -I (2000). Acacia tortitis trees in the woodlands had branches removed, were debarked, uprooted and broken. Acacia tortitis trees in the riverine had lower levels of utilization, whilst Acacia nilotica trees were mostly debarked. The method of elephant feeding varies within and between woody species, provided it is within the mechanical constraints of a certain size or species. Elephant behaviour is concluded to depend on spatiotemporal variation of forage abundance/quality, abundance of a preferred species, and species response (coppice or mortality). Elephant can cause a change of vegetation state, and increase spatial homogeneity of a plant population. The remnant population of woodland trees should provide the potential for recolonization, in which case the system would reflect the stable limit cycle. However, if browsing inhibits seedling recruitment, the system could reflect either a multiple stable state system or an artificial equilibrium imposed on a stable limit cycle. / Thesis (M.Sc.)-University of Natal, Pietermaritzburg, 2000.

Thesis (M.Sc. Agriculture (Pasture Science)) -- University of Limpopo, 2021 / Savanna rangelands are ecosystems which are characterized by the co-existence of scattered trees and shrubs with a continuous grass layer. However, the grass and tree balance has been highly altered as a result of disturbances caused by bush encroachment. Encroaching woody species have been shown to decrease species richness and abundance of the seed bank and ground‐layer diversity. So far little is known on the effect of bush encroachment and soil depth on the soil seed bank diversity in savanna rangelands. The objectives of this mini-dissertation were to (1) determine the influence of soil depth on soil seed bank diversity in bush encroached savanna rangelands, and (2) determine the relationships between soil seed bank herbaceous vegetation and physicochemical properties in encroached rangeland. In order to address these objectives, a savanna rangeland was demarcated into two encroachment gradients spanning from open to encroached rangeland. Within each encroachment gradient, six plots of 10 m x 10 m were randomly selected, whereby soil sampling and herbaceous vegetation were carried out and determined. In each replicate plot per encroachment level, five soil samples were randomly collected at 0-10 and 10-20 cm depths. The number of seedlings of different species emerging from the soil samples was used as a measure of the number of viable seeds in the soil and the composition of the seed bank using the seedling emergence method. The total seed densities showed significant differences (P<0.05) in the 0-10 cm depth layer in the open rangeland and encroached rangeland. Bush encroachment significantly (P<0.05) decreased the seed density of perennial grasses, specifically in 0-10 cm depth layer. Further, species diversity increased with bush encroachment in the 10-20 cm depth layer. Menhinick’s richness index showed no significant difference in the open and encroached rangeland, while species evenness decreased in the 0-10 cm depth layer and increased at 10-20 cm depth.The study also revealed negative correlations between organic carbon, calcium, clay, silt and forbs while mean weight diameter (MWD), a measure of soil aggregate stability was positively correlated with forbs. The canonical correspondence analysis (CCA) showed that pH, phosphorus, potassium and calcium were positively correlated to Eragrostis curvula and magnesium was negatively correlated to Panicum maximum. In open rangeland, CCA revealed that clay content was negatively correlated with species evenness while xii magnesium was negatively correlated to the Shannon Weiner index. Further, silt content was positively correlated with species richness and evenness. In the encroached rangeland, the CCA showed a negative correlation between magnesium and the Shannon Weiner index. The Sørensen’s index between soil seed banks and aboveground vegetation was low with index values of 0.22 and 0.24 in open and encroached rangeland, respectively. / AgriSeta

Thesis (M. Sc. Agriculture (Soil Science)) -- University of Limpopo, 2021 / Globally and in most parts of South Africa, there is a trend of increasing shrub encroachment in savanna rangelands. A number of studies have investigated the impact of shrub encroachment on soil organic carbon content (SOC) and soil organic carbon stocks (SOCs) in savannas. So far there is no clear consensus on whether shrub encroachment increases or decreases the level of SOC and SOCs, especially in semi arid savanna grasslands. Furthermore, knowledge on the effects of shrub encroachment on SOCs is largely restricted to the topsoil, as this is the part of the profile influenced by inputs and losses of soil organic matter. How shrub encroachment affects the vertical distribution of SOCs is rarely considered in the existing literature and the edaphic factors controlling SOCs with depth are poorly understood. The objectives of this study were (i) to quantify the vertical distribution of SOC and SOCs and (ii) to identify the edaphic factors controlling the vertical distribution of SOC and SOCs in a shrub-encroached savanna grassland sited on shallow plinthic soil. To achieve the objectives, a vegetation and soil survey of the savanna grassland was conducted whereby sampling areas were demarcated and characterized into open and shrub encroached grassland plots. In each encroachment level, three pits were randomly dug to the limiting layer on plots sited on the same soil type and similar topographic position. Soil samples were collected from the pits at depth intervals of 0-10, 10-20, 20-30, 30-40, 40-50, 50-60 and 60-70 cm. The collected soil samples were analyzed for chemical and physical properties in the laboratory. Correlation analysis was carried out to determine the relationship between SOC and SOCs, which were the variables of interest in this study and related controlling soil physicochemical properties. The results showed that SOC was significantly greater (P<0.05) in the shrub-encroached grassland compared to open grassland. Furthermore, the results revealed that SOC was on average 19 and 13% greater in the topsoil (0-20 cm) and subsoil (20-70 cm) of shrub encroached grassland compared to open grassland. The greater SOC in the topsoil of the shrub-encroached grassland was mainly attributed to higher SOC inputs from plant litter and detritus derived from trees and grasses, which are the dominant plant life forms in savannas. In the topsoil, SOC and SOCs were positively correlated with extractable phosphorus (P) (r = 0.60; P < 0.05), while in the subsoil they were positively correlated ix with extractable phosphorus (r = 0.54), soil porosity (r = 0.52), extractable copper (r = 0.46), extractable zinc (r = 0.41), exchangeable calcium (r = 0.37) and negatively correlated with mean weight diameter (r = -0.43). Overall, SOC and SOCs under both shrub-encroached and open grasslands vertically decreased with soil depth. The results obtained highlighted that the factors controlling the level of SOC and SOCs differs in the topsoil and subsoil of the studied shrub encroached grassland. These findings suggest that in the shallow plinthic soil investigated in this study, SOC in the topsoil is controlled by the macronutrient P, while in the subsoil it is physically protected by soil aggregates and chemically stabilized via complexation interactions with exchangeable cations and heavy metals. In-depth understanding of the physico-chemical factors controlling SOC storage is critical to foster management practices that will improve the cycling of SOC in shrub-encroached savanna grasslands.

A major challenge in savanna rangeland studies is estimating woody vegetation cover and densities over large areas where field based census alone is impractical. It is therefore crucial that the management and conservation oriented research in savannas identify data sources that provides quick, timely and economical means to obtain information on vegetation cover. Satellite remote sensing can provide such information. Remote sensing investigations, however, require establishing statistical relationships between field and remotely sensed data. Usually regression is the empirical method applied to field and remotely sensed data for the spatial estimation of woody vegetation variables. Geostatistical techniques, which take spatial autocorrelation of variables into consideration, have rarely been used for this purpose. We investigated the possibility of improving woody biomass predictions in tropical savannas using cokriging. Cokriging was used to evaluate the cross-correlated information between SPOT (Satellites Pour l’Observation de la Terre or Earth-observing Satellites)-derived vegetation variables and field sampled woody vegetation percentage canopy cover and density. The main focus was to estimate woody density and map the distribution of woody cover in an African savanna environment. In order to select the best SPOT-derived vegetation variable that best correlate with field sampled woody variables, several spectral vegetation and texture indices were evaluated. Next, variogram models were developed: one for woody canopy cover and density, one for the best SPOT-derived vegetation variable, and a crossvariogram between woody variables and best SPOT-derived data. These variograms were then used in cokriging to estimate woody density and map its spatial distribution. Results obtained indicate that through cokriging, the estimation accuracy can be improved compared to ordinary kriging and stepwise linear regression. Cokriging therefore provided a method to combine field and remotely sensed data to accurately estimate woody cover variables. / Thesis (M.Sc.)-University of KwaZulu-Natal, Pietermaritzburg, 2008.

Balancing increasing elephant numbers with biodiversity conservation in small reserves has become a concern for many protected area managers. Elephants are considered important agents of disturbance creating heterogeneity and thus contributing to the maintenance of biodiversity. However elephants also damage vegetation through their destructive feeding habits, and this has led to pressure to reduce elephant populations in many reserves. Quantitative data on the impact of elephants on invertebrates, the main component of biodiversity at the species level, are lacking. The aim of this project was to assess the effect that habitat alteration by elephants has on the diversity of selected ground-dwelling invertebrates (ants, centipedes, millipedes, spiders, scorpions and termites) through the provision of logs and dung as a potential refuge niche for these invertebrate communities, and to determine the effect of spatial (vegetation types) and temporal (season and age of dung) variation on the invertebrates using these refugia. Variation in impacts was considered important because savanna is not homogenous and the impact of the refugia is likely to be dynamic in terms of seasonal trends in invertebrate populations, and in terms of changes in the environmental conditions offered by the refugia. Elephant impact on vegetation, quantity of refugia (logs and dung) produced and invertebrate diversity associated with refugia were determined for 115 transects within Madikwe Game Reserve in the North Western Province, South Africa. Invertebrate abundance, species richness and diversity were always higher under refugia than in areas without refugia. Vegetation utilisation, frequency of refugia production and invertebrate diversity showed strong temporal variation (seasonal); elephant impact and production of logs were higher in winter than in summer because elephants are more likely to feed on woody vegetation in winter when grass nutrient levels are low. Invertebrate diversity under the logs was higher in summer than in winter, and this probably reflected the higher abundance and diversity of invertebrates that are usually associated with the warmer, wetter summer months. The effect of adding refugia to three vegetation types on invertebrate diversity was tested experimentally at Makalali Private Game Reserve in the Limpopo Province, South Africa. Logs and elephant dung were set out in five plots each measuring 20m x 20m within Govender - iii mixed bushveld, riverine and mopane woodland. Significant differences were observed in invertebrate abundance, species richness and diversity between the refugia and control plots that lacked refugia and between the three vegetation types sampled. Similarity between invertebrate communities utilising the different refugia types and between the three different vegetation types were tested using the Jaccard similarity coefficient. The three vegetation types shared fewer than 50% of their species, as did the logs, dung and control sites. However the results obtained do illustrate a higher degree of similarity between the refugia substrates (logs and dung) than the control sites and between the more heterogeneous vegetation types (mixed bushveld and riverine) than the mopane veld. This indicated that invertebrate communities associated with refugia were not uniform, but were influenced by vegetation type. An experimental test of temporal changes in invertebrate community composition illustrated the importance of elephant dung as a microhabitat for different invertebrate groups over different ages of dung (three days, two, four, 12 and 32 weeks old). Colonisation of the dung, by dung beetles was immediate but as the microclimate of the dung changed with time, the new conditions were ideal for other invertebrate taxa. Over a period of eight months, the change of invertebrate communities utilising the dung included dung beetles, followed by millipedes and [mally ant and termite communities. The results of this study illustrated the importance of refugia (logs and dung) produced by elephants for ground-dwelling invertebrate species in the savanna environment. The extent of the influence of the refugia varied both spatially and temporally and this should be considered in future monitoring or in measuring impacts. While further research on a broader range of organisms and at larger scales is necessary, elephants do have a positive impact on at least some components of biodiversity, through the process of facilitation of refugia. / Thesis (M.Sc.)-University of KwaZulu-Natal, Pietermaritzburg, 2005.

Browse selection, intake, utilisation, palatability, quality and production are tightly linked and need to be considered together in trying to improve our understanding of browsing dynamics and the interactions between browsers and vegetation. Such an understanding is necessary in order to re-evaluate determinations of browser carrying capacities and evaluating actual and potential impacts of browsing animals on vegetation composition and diversity. Browser carrying capacity is determined by both the quantity and the nutritive value of forage. The measurement of browse quantity and nutritive value and the matching of browse supply to browser demand are central to sustainable utilisation and the monitoring of vegetation health. South African savannas are poorly studied with respect to tree canopy growth and browse production making it difficult to quantify the available browse biomass on which browsing capacity estimations are based, and consequently difficult to estimate levels of browsing that are sustainable. This study addressed these issues by investigating browse dynamics, broadly aiming to (1) explore factors affecting browse production, biomass and nutritive value; (2) develop models to assess and monitor these parameters across seasons and properties; (3) use the resultant models in improving our understanding of how to determine browser carrying capacities. More specifically, our study sought to examine the effects of plant physiognomy, forage nutritive value, canopy stratum, defoliation, temperature, rainfall and soil nutrient status on the browse production of evergreen (Carissa bispinosa, Euclea divinorum, Gymnosporia senegalensis), semi-deciduous (Spirostachys africana, Ziziphus mucronata) and deciduous (Acacia nilotica, Dichrostachys cinerea) savanna tree species from June 2003 – June 2005 in three sites along the northern Zululand coastline of KwaZulu-Natal. Available browse biomass, during the dry season, of four key savanna tree species (A. nilotica, E. divinorum, G. senegalensis, and S. africana) was estimated through the development of allometric regression equations. Non-linear regression was used to investigate the relation between the leaf dry mass (LDM) and canopy volume (CVol) of each of the four tree species. Exponential regression (y = a + brlnx) of the natural logarithm of CVol data provided the most accurate and precise description of the tree CVol – LDM relation. A study was undertaken to determine which factors may influence browse production in a southern African savanna. Regression tree models for the browse production identified that the dominant factors influencing browse production were CVol (m3), season, species and height to the lowest leaves of the tree canopy (HL) (m). The length of the growing season had a marked effect on the production potential of savanna tree species, suggesting that improved conditions for growth, i.e. greater rainfall, soil moisture content and improved soil nutrient availability result in a longer period of rapid sustained growth. Species was identified as an important contributing factor to differences in browse production rates, suggesting the need for the development of species or species group models. Mean annual browse production of evergreen trees was greater than that of deciduous and semi-deciduous trees. Mean quarterly (three monthly) browse production was highest, for all trees, during the wet season, with the greatest difference between wet and dry season production being observed in deciduous forms. Evergreen forms showed continuous growth over the whole study, with enhanced growth over the wet season. Deciduous forms, on the other hand, concentrated growth in spurts, when environmental conditions became favourable, with most production occurring during a short growing season. Browse nutritive value was found to be greatest during the wet season, when growth and photosynthesis are at their greatest. Further, browse nutritive value was greatest in deciduous species. Evergreen trees were found to have greater acid detergent fibre (ADF) concentrations than both the deciduous and semi-deciduous trees. By contrast, crude protein (CP) concentrations were greater in semi-deciduous and deciduous species than in evergreen species. The daily CP requirements for maintenance for an adult impala (45 kg) were met by all species over all three study areas and all seasons. Daily CP requirements for growth and lactation, however, were only ever met by deciduous and semi-deciduous species, though this result was not consistent over study areas and seasons. Predictive models for the production of browse on deciduous, semi-deciduous and evergreen trees in northern Zululand were developed using multivariate adaptive regression spline functions. The best predictors of growing season browse production in all three tree guilds (defined here as a group of trees having a characteristic mode of living) were primarily measurable tree dimensions, while the prevailing environmental conditions had little impact. Differences in the production, nutritive value and available browse biomass between the different tree forms and seasons have a profound effect on the determination of browser carrying capacities and need to be incorporated into any game or conservation management plan. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2012.

The effects of fire and mechanical clearing were investigated for their potential in assisting with the eradication of Chromolaena odorata (previously Eupatorium odoratum). The study was divided into two focus areas, the first focused on mechanical clearing of dense stands of C. odorata on three sites and the second focused on the long term influences of a single burn on C. odorata plants in the different size categories. For mechanical clearing, two key issues were investigated; namely whether this type of clearing procedure was effective in dense C. odorata stands and whether rehabilitation was necessary in these cleared areas. The study was conducted from July 2002 to June 2004. The area was subject to a severe drought throughout the duration of the study. The severe drought had a large influence on the result in both focus areas. A bulldozer was found to be a very effective at clearing dense C. odorata stands. Results from the mechanical clearing study showed that there was still a large viable grass seed population in the areas that had been covered by a dense stand of C. odorata plants for over three years. Thus, indigenous plants were able to re-colonize the area after removal of C. odorata without human intervention, even thought the area was experiencing a severe drought. The density of C. odorata seedlings emerging in the cleared areas was far lower than expected. The C. odorata density in the permanent plots, for seedlings that germinated in the first season after clearing (SeptemberOctober 2002), was only 0.25,0.03 and 0.72 per 5 m2 in the three sites respectively by the end of the study in June 2004. For the C. odorata seedlings that germinated in the second season (September-October 2003) the density was, 0.5, 0.56 and 1.06 per 5 m2 in the three sites respectively by the end of the study in June 2004. It was suspected that the drought influenced seed germination. Unfortunately the number of C. odorata seedlings was so low, that no significant relationship could be found between grass and C. odorata seedling density. By the end of the study the grass fuel mass in all the rehabilitated sites was already over 3000 kg ha-1, even though the area was experiencing a severe drought. This grass fuel load, when burnt, will assist land managers in controlling C. odorata plants, especially seedlings. Very few other alien invasive plant species emerged in the cleared areas. At the Mhlosinga site, Senna pendula made up less than one percent of the herbaceous species composition and only a single Ricinus communis plant was recorded. No alien plant species were recorded on the other two sites. Results from the burning trials revealed that plants in all the size categories were affected by fire. Greater fuel masses and fire intensities were required to kill larger C. odorata plants relative to smaller ones. Fire was found to be very effective at eliminating small and medium size C. odorata plants. Fire applied as a once off treatment had a significant long-term effect on the C. odorata population. The following fuel loads were required to achieve 80% mortality in this 11 study: for small plants a fuel load of over 4000 kg ha-I, for medium plants a fuel load over 4200 kg ha-I and for large plants a fuel load over 4600 kg ha-I. Little difference could be detected between a head or a back burn, as both fire types had their own advantages and disadvantages. Although some of the C. odorata plants in the burnt plots had not perished by the time of the first investigation, following the burn (February 2003), by the time of the second investigation (June 2004), many of these plants had eventually succumbed. These results highlighted the fact that plants which are damaged by fire were more likely to persish during an extended droughts period, than plants which were not subjected to fire. Results from the control plots, in the burning trials, for medium and large plants showed dramatic increases in density over time. Tagged individuals from the control plots did reveal that some of the medium and large plants did die during the drought, although the amount was negligible when compared to the number of new plants growing into the new size categories. A large proportion of the small plants in the control plots also survived the drought with many of them even growing into the medium category. The difference between the control plots and the burnt plots was obvious and significant, especially once the fuel mass exceeded 3783 kg ha-I. Results from this study show that fire can be used as a very effective tool in assisting land managers to control C. odorata in open savanna bushveld. / Thesis (M.Sc.)-University of KwaZulu-Natal, Pietermaritzburg, 2006.