A dissertation submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfilment of requirements for the degree of Master of Science. August 2015. / iii ABSTRACT Global changes are likely to have negative impacts on many ecosystems including savannas. Semi-arid environments are notable for the wide range of seasonal patterns of leaf display in the tree communities. The environmental cues of leaf out and leaf drop are not consistent across species, and are not always directly linked to water availability, indicating that some species might be particularly sensitive to changes in climate. Strategies employed by trees which leaf early or drop their leaves late are likely to impact other aspects of their life-history and functioning so I expect particular plant functional types to be associated with particular vegetation functional traits. I assessed how variable savanna leafing strategies are among 28 species at a semi-arid savanna site at Nylsvley, and used this information to group species into plant functional types (PFTs). These PFTs were then assessed in terms of key vegetative traits to explore the life history consequences of different leafing strategies. Leaf phenology was monitored throughout one growing season and quantified using 8 key phenological metrics. The timing of leaf display tracks the timing of seasonal rainfall but with wide variation, with some species retaining their leaves throughout dry season. Other species loss some leaves throughout the growing season, some species only flushed their leaves after the first rains, and other flush before the first rains. I identified 4 clear PFTs using the MClust clustering integrated with subjective procedure. Four vegetative traits were measured: specific leaf area, leaf nitrogen, maximum stomatal conductance and wood density. I identified some clear trade-offs between vegetative traits and phenological strategies. There was also a positive relationship between degree of rain stimulated flushing metric and wood density. Using objective clustering methods to determine plant functional types has some clear advantages over more subjective methods but depends on good input data. Identifying plant functional types at Nylsvley has led to some insights into functioning of these savannas, but as there appear to be strong links between plant traits and particular leafing strategies it might be more appropriate to explore syndromes of vegetation functional traits when modelling responses to global change.

The thesis presents a thorough, in-depth study that fills some of the gaps in the knowledge of the impacts of woodland utilisation in communal areas. The chosen case study villages are in Bushbuckridge, a government gazetted Integrated Sustainable Rural Development programme node, making the results pertinent to sustainable energy policy reform in South Africa. A case-study of two villages was used to investigate the spatial and structural changes in fuelwood supply in response to fuelwood extraction as well as the changes in use-patterns over time. A survey of the structure and composition of the woody vegetation and wood harvesting patterns around the villages was conducted and compared against historical data, spanning 17 years. Total wood stock in the communal woodlands of both villages declined over the study period; the loss being greater in Welverdiend. Significant, negative change in the structure and species composition, particularly of species that are commonly harvested for fuelwood has occurred in Welverdiend but not in Athol. The absence of negative impacts in Athol implies that harvesting regimes here are more sustainable but it is more likely that this is due to the lower human population and lower fuelwood extraction pressure. The changes in woodland structure were linked to landcover change patterns that occurred in the villages over the last 44 years, from their creation through forced resettlements on old farms in the area. Landcover change patterns were similar in both villages since 1965 but there was significantly greater woodland loss in Welverdiend (48% woodland loss) in comparison to Athol (25% woodland loss). The systematic loss of woodland areas to agricultural fields was linked to expanding residential areas due to human population growth. Deforestation occurred where woodlands were already impacted through selective harvesting. The physical changes in woodland structure and landcover were linked to a detailed socio-economic analysis of the two villages, providing critically important data for the sustainable management of woodlands in South Africa. The impact of access to electricity on fuelwood consumption rates was carried out through analysis of the economic, time and opportunity costs of fuelwood collection, compared against the different fuelwood availability in each village. In Welverdiend demand for fuelwood has so far proved inelastic; households have adjusted their fuelwood collection regimes, going on fewer collection trips but spending longer times for each trip but ultimately household investment is similar to that in Athol. Fuelwood demand is maintained in Welverdiend by the availability of purchased fuelwood and harvesting in new sites. A model to predict the socio-economic factors at the household and per capita level which affect fuelwood consumption was developed. Revealing in the process that households with access to electricity used less fuelwood annually and the amounts of fuelwood used were influenced by the household perceptions of fuelwood scarcity in the village, Household population size had a direct bearing on the likelihood of households switching to electricity with every addition to the household size decreasing the likelihood of switching by 48%. This study has major implications for the government’s on-going rural electrification programme. Interventions are required that raise awareness about fuelwood availability trends, based on landscape developments and targeting women as the main users of fuelwood.

Thesis (Ph. D.)--University of Texas at Austin, 2004. / Supervisor: Norma Fowler. Vita. Includes bibliographical references.

Thesis (doctoral)--Swedish University of Agricultural Sciences, 2003. / Appendix includes reprints of five manuscripts, four co-authored with others. Includes bibliographical references. Also partially issued electronically via World Wide Web in PDF format; online version lacks appendix.

A dissertation submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfilment of the requirements for the degree of Master of Science Johannesburg, September 2015 / Plant functional traits provide a means to investigate the diverse ecological strategies employed by plants and a tangible link to assess how the variability in these traits might influence ecosystem processes and functioning. The aim of this dissertation has been to determine how plant and soil nutrient dynamics in a savanna environment are affected by two primary drivers, one a top-down driver, being herbivory by large mammalian herbivores and the other a bottom-up driver, the variable N2-fixation capacity of tree species. To the best of my knowledge this is the most comprehensive study to date to investigate the bioavailability of soil nutrients and the link between these availabilities and plant functional traits. Furthermore this study provides important insight into the use of a novel technology, ion exchange resin capsules in a South African savanna context. By studying a selection of plant functional traits (nutrient concentrations, ratios as well as specific leaf area, relative chlorophyll content and leaf dry matter content) and soil nutrients (suite of macro- and micronutrients) associated with two species of savanna tree of contrasting N2-fixation capacities, I went about investigating how herbivory differentially influences the nutrient dynamics of this system. Selecting individuals of the N2-fixing Acacia tortilis and the non-N2-fixing Combretum hereroense both inside an exclosure and on the adjacent land allowed me to determine the potential impacts by herbivores. These include both direct impacts from foraging and indirect impacts through the regulation of nutrient input pathways via deposition of dung and urine. The work compiled for this dissertation is based on the experimental work conducted in a mesic savanna system in the Marakele Park (PTY) Ltd. During the course of this dissertation, I investigated herbaceous and woody biomass in relation to protection from and exposure to herbivory, determining any differences in the functional leaf traits between individuals inside and outside the exclosure, if these differences were exhibited in the associated herbaceous biomass as well as a comprehensive assessment of the bioavailability of 15 important micro- and macronutrients using ion exchange resin capsules. These capsules were incubated in the soil over the entire summer rainfall period, providing a cumulative view of nutrient bioavailability during the growing season. In this work I also demonstrated whether particular nutrients are associated with specific drivers (i.e. herbivory, canopy position or N2-fixation). Furthermore, these results were then looked at together to suggest the mechanism by which herbivory and N2-fixation drive nutrient dynamics and make recommendations on the use of these results in managing savanna systems in the future. Between the two sites, aboveground herbaceous biomass was significantly greater when protected from herbivores than on the adjacent land. Both exposure to herbivory and N2-fixation capacity were found to alter plant functional traits. Herbivore presence was associated with an increase in herbivore-resistant or structural traits such as C/N, C/P, foliar C and SLA as well as a reduction in N and P content. These less palatable leaves were accompanied by a significantly lower availability of a number of important soil elements, namely NO3-N, inorganic N, P, K, Na, Cu, B, Mg, and S. This suggests a feedback loop between these two components of the ecosystem. N2-fixation capacity is associated with greater concentrations of elements such as N and P and a reduction in traits that are illustrative of a greater structural investment into leaves. Soil nutrient bioavailability however, shows a reduction in certain nutrients when associated with Acacia. A number of nutrients which show a reduction in availability are those which are essential to N2-fixation machinery, namely B and Fe but also lower bioavailabilities of Al and Mg. Finally, Ca, NO3-N, B, Fe, Al and inorganic N were found in greater quantities below the tree canopy than beyond it. In conclusion both herbivory by large mammalian herbivores and N2-fixation have significant effects on tree health, through their regulation of limiting nutrients and alteration of leaf traits. Given the changes which these drivers are capable of exerting on plant and soil nutrient dynamics, this has important consequences for ecosystem processes and functioning and highlights potential considerations in the long-term sustainable management of savannas.

A dissertation submitted to the Faculty of Science, University of the Witwatersrand, in partial fulfilment of the requirements for the degree of Master of Science, Johannesburg, South Africa 2017. / Fuelwood is still heavily relied upon by rural communities as a source of energy. The current levels of wood harvesting have been deemed unsustainable, with models predicting the local exhaustion of wood resources in most cases. However, wood depletion has generally not happened to the level of severity predicted by the models. This may partially due to under-accounting for coppice regeneration. Many savanna species that are harvested for fuelwood demonstrate strong coppicing ability, which allows for regrowth after a disturbance. This ability to regrow or coppice is a key functional trait which allows species to persist and survive in frequently disturbed environments. There is surprisingly limited knowledge about coppice dynamics in savanna trees and how managerial actions might influence the coppice response and production of savanna species. To address this problem, this study investigated the influence of four post-harvest treatments on various aspects of the coppice response of Terminalia sericea and Dichrostachys cinerea – two important savanna fuelwood species – in a field experiment. A total of 108 felled trees per species were exposed to one of four treatments, which were applied monthly for 12 months. The treatments were (1) Control – no coppice shoots were removed for 12 months, (2) Harvest – all coppice shots were removed monthly, (3) Single prune – the coppice shoot with the widest diameter was left on the stump, and (4) Usable – coppice shoots that reached a diameter of 2 cm were removed. The effect that these treatments had on the average diameter, length and cumulative number of coppice shoots produced per stump was compared. The measurements of diameter and length were used in developing allometric equations for the prediction of coppice shoot biomass. The predicted biomass produced through the study was compared across treatments to gain an understanding of how productivity could be influenced by management of coppice shoots. The mean shoot diameter, length, as well as cumulative number of coppice shoots produced per stump was higher in the single prune treatment for both species while the harvest treatment resulted in high numbers of coppice shoots but with low average diameter and length. Applying the growth rates of coppice shoots found in this study it can be assumed that unmanaged coppice shoots will take approximately 5.5 years to reach a preferred harvestable diameter of 4 cm, while single prune coppice shoots would take 3.3 years to reach the preferred harvestable diameter. Diameter had more of an influence on the predicted coppice biomass production of T. sericea while D. cinerea biomass prediction was more influenced by shoot length. However, only diameter was used to compare the prediction of coppice biomass with previously developed equations as these equations did not consider length for predicting biomass of different components of trees. From the derived models, the calculated biomass at the end of the study period as well as the calculated biomass produced through the year was greatest for the control treatment. The predicted dry shoot stem biomass at the harvestable diameter of 4 cm was 114.60 g for T. sericea and 95.88 g for D. cinerea. From the findings of this study it is clear that post-harvest management can be utilised to manipulate coppice response and biomass production. Keywords: Fuelwood; Coppice shoot; Post-harvest treatment; Diameter; Length; Biomass / LG2018

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The growth and recovery of trees from disturbances such as fire and browsing is driven by the intensity of the disturbances and the availability of resources. In savannas, resprouting has become recognized as a key functional trait in plant ecology over the past decade. Although this may indeed be the case, there is still limited information about the physiology and growth strategies of resprouting trees. Available information about the influence of disturbance comes from ecosystems that are in many ways different from tropical and sub-tropical savannas. Therefore it is important to know and understand post-disturbance tree responses and limitations so as to establish sustainable use and management practices. This thesis reports the findings of a study, conducted in the Mpumalanga province of South Africa, aimed at achieving a better understanding of the influences of resource availability (water and nutrients) and disturbances (herbivory and repeated cutting) on the coppicing of a widely distributed savanna tree species that is both ecologically and economically important; Terminalia sericea. To investigate the effects of resource availability, cut trees were exposed to different levels of water and nutrient (nitrogen and phosphorus) supplementation over a period of two years in a factorial experimental design. A number of coppice regrowth variables (e.g. shoot production, resprout shoot diameter and shoot length) were measured monthly, while the phenological responses (e.g. timing of leaf discolouration and fruit presence) were monitored every two weeks. The effects of disturbances were investigated in two separate experiments, in which cut trees were exposed to a five month browsing period and different cutting frequencies, respectively. Coppice regrowth variables were monitored for 12 months in trees exposed to browsing and for five months in trees exposed to different cutting frequencies. The effect of multiple cutting cycles on total non-structural carbohydrates and leaf chemistry (carbon, nitrogen and phosphorus) was measured. There was evidence of self-thinning of coppice shoots within cut trees in all experiments in the second year of growth, with supplemented trees also recording lower shoot numbers. With shoot production higher in trees that received no resource addition, water and nutrients interacted synergistically, doubling shoot diameter and shoot length for supplemented trees after 12-months of addition. The majority of the phenophases monitored peaked in the wet growing season. Supplemented trees changed colour from the typical green to senescent yellow later in the growing season than unsupplemented trees. Fruit presence occurred in the second year after cutting for supplemented trees. Shoot length and shoot diameter for unbrowsed trees were twice those for browsed trees, with leaf nitrogen and phosphorus content significantly higher for browsed trees compared to unbrowsed trees. Trees subjected to multiple cutting cycles recorded half the TNC levels, and half the resprout shoot diameter and shoot length of trees cut only once. Results from this study demonstrate that self-thinning (i.e., negative change in shoot number) is not primarily under resource control. By contrast, the findings suggest that shoot growth characteristics, the timing and duration of phenological stages in coppicing trees are resource-limited in savannas. Browsing induces an initial compensatory response through higher shoot production in browsed trees and should be kept minimal because, in the long run, browsed cut trees would take longer to recover lost biomass compared to unbrowsed cut trees. Repeated cutting significantly depleted non-structural carbohydrate reserves in stems, implying that repeatedly cut trees rely heavily on non-structural carbohydrate reserves for regrowth. The significantly lower shoot diameter and shoot length in repeatedly cut trees imply that the coppicing ability of a tree reduces as the cutting frequencies increase. Such information can be vital in establishing the competitive growth ability of T. sericea in a multiple-species ecosystem affected by changes in resource availability as well as natural disturbances. With reference to multiple cut trees, a form of cutting or harvesting strategy should be in place that allows for sustainable regeneration of the study species. A threshold in terms of number of cutting cycles a tree can tolerate based on either maximum or minimum levels of reserve carbohydrates should also be established, as this has a direct effect on the coppice growth and survival.

Thesis (Ph. D.)--University of Pretoria, 2005. / Title from PDF t.p. (viewed Mar. 27, 2006). Includes bibliographical references.

Thesis (MScFor)--Stellenbosch University, 2002. / ENGLISH ABSTRACT: Increases in woody plant density ('bush encroachment') reduce livestock production and biodiversity. By convention, soil moisture, soil nutrients, fire and herbivory are regarded as the principal factors governing the tree-grass ratio of savannas. An experiment with a completely-crossed design was employed to investigate woody seedling (Acacia me/lifera) recruitment near Kimberley, Northern Cape, South Africa, after fire and under conditions of maximum-recorded rainfall, nitrogen addition and grazing. The field experiment was repeated as a garden experiment to determine if the two experiments yield comparable results. Tree germination in the field was extremely low, probably due to below-average natural rainfall in plots that only received natural rain, and insufficient watering frequency in irrigated plots. As a result of low germination, none of the treatments (rain/nitrogen/fire/grazing) had a significant effect on tree recruitment in the field experiment. The duration of the experiment (2000/2001 growing season) was insufficient for the treatments to affect grass composition, although the high rainfall treatment and grazing exclusion significantly improved grass cover and height. The garden experiment showed that frequent watering, no nitrogen addition and grass clipping significantly enhanced tree germination and survival (termed 'recruitment'). There were also significant interactions among rain, nitrogen and grazing in their effects on tree recruitment. The effects of rain on tree recruitment were more pronounced under nitrogen supplementation and vice versa. Similarly, high rain and high nitrogen enhanced the effect of grazing on tree recruitment. It is inferred that above-average rainfall years with frequent rainfall events are required for mass tree recruitment. Tree seedlings can further benefit from space and resources which are made available through grass defoliation. Conversely, nitrogen enrichment improves the competitive ability of the fast-growing grasses relatively more than that of the N2-fixing tree component, thereby suppressing tree recruitment. In contrast to conventional wisdom that grazing alone causes encroachment, it is suggested that there are complex interactions between the abovementioned factors and 'triggering' events such as unusually high rainfall. Contrary to many claims that equilibrium models are inappropriate for explaining savanna dynamics, it was shown that consumer-resource theory has explanatory power for bush-grass dynamics of the savanna studied. The state-space approach that was used facilitated the understanding of savanna dynamics and enabled predictions about the system's response to perturbations. The applicability of consumer-resource theory to semi-arid nutrient-poor savannas confirmed the importance of resource competition in structuring natural systems. / AFRIKAANSE OPSOMMING: Toenames in die digtheid van houtagtige plante ('bosverdigting') verlaag veeproduksie en biodiversiteit. Volgens konvensie word grondvog, grondvoedingstowwe, vuur en beweiding beskou as die belangrikste faktore wat die boom-gras verhouding van savannas bepaal. 'n Volledig-gekruisde ("completely crossed") eksperiment is ontwerp om boomsaad-ontkieming (Acacia mellifera) naby Kimberley, Noord-Kaap, Suid- Afrika, te bestudeer na 'n brand en onder toestande van maksimum-aangetekende reënval, stikstof toevoeging en beweiding. Die veldeskperiment is herhaal as 'n tuineksperiment om vas te stel hoe resultate van die twee eksperimente vergelyk. Boomsaad-ontkieming was uiters laag in die veld, waarskynlik weens ondergemiddelde reënval in persele wat slegs natuurlike reënvalontvang het, en 'n te lae benattingsfrekwensie in die besproeide persele. As gevolg van lae ontkieming in die veldeksperiment, het geen behandeling (reënval/stikstof/brand/beweiding) boomsaad-ontkieming beduidend geaffekteer nie. Die duur van die eksperiment (2000/2001-groeiseisoen) was te kort vir die behandelings om grassamestelling te beïnvloed, alhoewel besproeiing en geen beweiding die grasbedekking en -hoogte betekenisvol verhoog het. Die tuineksperiment het getoon dat boomsaadontkieming en vroeë oorlewing (genoem 'vestiging') betekenisvol verhoog is deur gereëlde benatting, geen stikstof toevoeging en die sny van gras. Daar was ook beduidende interaksies tussen reën, stikstof en beweiding in hul invloed op boomvestiging. Die reënbehandeling het 'n groter uitwerking op boomvestiging gehad onder stikstoftoevoeging en vice versa. Hoë reënval en stikstoftoevoeging het ook die invloed van beweiding op boomvestiging versterk. Die afleiding is dat bogemiddelde reënvaljare met gereëlde reënbuie 'n vereiste is vir grootskaalse boomvestiging. Boomsaailinge kan verder voordeel trek uit die spasie en hulpbronne wat beskikbaar raak wanneer gras ontblaar word. Daarteenoor verhoog stikstofverryking die mededingendheid van die vinniggroeiende grasse meer as dié van die N2-bindende boomkomponent, met die gevolg dat boomvestiging onderdruk word. In kontras met die konvensionele veronderstelling dat beweiding opsigself bosverdigting veroorsaak, word voorgestel dat ingewikkelde interaksies plaasvind tussen die bogenoemde faktore en ander 'sneller-gebeurtenisse', soos buitengewoon hoë reënval. In teenstelling met baie aansprake dat ekwilibrium modelle ontoepaslik is om savanna-dinamika te verklaar, is getoon dat die verbruikers-hulpbron teorie ("consumer-resource theory") oor verklaringsvermoë beskik vir boom-gras dinamika van die savanna wat bestudeer is. Die staat-spasie ("state-space") benadering gebruik, het begrip van savanna-dinamika bevorder en voorspellings moontlik gemaak aangaande die sisteem se reaksie op versteurings. Die toepaslikheid van verbruikers-hulpbron teorie vir semi-ariede, nutriënt-arm savannas bevestig dat kompetisie vir hulpbronne 'n sentrale rol speel in die strukturering van ekologiese sisteme.