Thesis presented for the Degree of Doctor of Philosophy at the University of the Witwatersrand / Successional changes in the vegetation after fire were studied in several fynbos communities of the south-western Cape Province of South Africa. The study sites were located in the mountains, at altitudes between 300 and 1000 m a.s.l., in areas with winter rainfall regimes -1 and annual precipitation of about 900 to 1000 mm. yr Soils are highly leached, derived principally from quartzites. The two main sites were Zachariashoek near Paarl, where summers tend to be rather dry, and Jakkalsrivier east of Grabouw, where summer drought is ameliorated by fog precipitation and cloudiness. Successional changes were followed for intervals of up to 10 yr between fires, as well as for similar periods in vegetation that had been unburnt for 25 yr. Vegetational changes were analysed by means of repeated floristic assessments on permanent quadrats and point-quadrat sampling of canopy cover composition on these and on larger plots. At Jakkalsrivier, recently burnt and long unburnt vegetation were also compared by paired samples. Demographic trends in populations of prominent shrub species were followed by repeated censuses of tagged samples in unburnt and recently burnt vegetation. Also at Jakkalsrivier, the effects of fire on resources available to plants were examined by sampling soil moisture and soil mineral nutrients, as well as by following trends in xylem pressure potentials in selected species of plants and analysing their foliar nutrient concentrations. Effects of fire on microclimate were tested by comparative studies on burnt and unburnt sites. All fynbos communities sampled proved to be highly stable in the face of fire. Essentially, the pre-fire species composition was regained in 2-3 yr in every case. Species were added after fire, partly because of the appearance of ephemerals with life histories tied to fire, but also because of the reappearance of longer-lived plants as well as through the readier detection of species in vigorous vegetative form. The species richness of the regenerating corrununities tended to be quadratically related to pre-fire biomass, as predicted from current succession theory. Most species in any corrununity (about 70% on average) regenerated vegetatively by sprouting after fire. The relative numbers of species that regenerated germinatively, i.e. the seeders, did not vary in a manner predictively related to corrununity biomass. There were relatively few species with specialised life histories based on reseeding, such as those with canopy-stored seed and ephemerals with presumably specialised requirements for germination. Virtually no recruitment could be found among plants in the older (about 25 yr) vegetation, in contrast with lowland fynbos sites, where recruitment of herbaceous species occurs, and some mountain fynbos sites on more fertile soils, where forest precursors may sometimes colonise. Canopy redevelopment after fire indicated similar resilience among the different corrununities, despite variation in regrowth rates. Pre-fire growth-form composition was restored within around 10 yr. Maximum leaf-area indices ranged from about 1,5 to 2,5, although corrununi ties on phreatic sites had leaf-area indices exceeding 3,0. There was no evidence for a suppression of the understoreys by overstorey layers, mainly because the latter were sparse despite the abundance of tall broad-sclerophyllous shrubs in certain habitats. This was because the taller shrubs had sparse or slender crowns, or both, and because mortality tended to thin the populations before dense canopies developed. Trends in the composition of the canopy varied among corrununities. corrununities in productive habitats, i.e. in this case on phreatic sites, were dominated in the early stages by a relatively luxurious growth of ephemeral herbs and soft shrubs which declined within around 3-4 yr. Other sites had very sparse ephemeral cover, the early stages being dominated mainly by Restionaceae, Cyperaceae, and other sprouting herbs, and sprouting and seeding shrubs, which were constituents of the pre-fire canopies. In this respect, the fynbos is clearly distinguished from the California chaparral, for example, where ephemerals tend to dominate the post-fire stages on most sites. There was no evidence that fire had any effect on the water relations of regenerating vegetation, although stream discharge is known to be increased by fire in these environments. There was tentative evidence, in enhanced foliar concentrations of some mineral nutrients, that regenerating species of climax plants exploited nutrients released in fire. However, any such responses were small, especially in comparsion with responses observed in chaparral, for example. Ephemeral shrubs had much higher concentrations of foliar nutrients overall than climax species, tending to confirm the correlations found in Australian heathlands between plant life-history and nutrient economy. The effects of fire on microclimate were pronounced, especially on the thermal and water vapour regimes experienced by seedlings and sprouts. These extremes did not, however, appear as water stress in regenerating plants. Despite relatively sparse canopies, mature vegetation did reduce light at the ground to levels likely to affect seedling recruitment and survival. Preliminary experiments with a local dominant shrub, Leucadendron xanthoconus, showed a pronounced intolerance of shading and hence that light attenuation by canopies must be implicated in successional processes. The demographic studies indicated that density-dependent effects were not important in survival of plants. Two species of fire ephemeral shrubs effectively died out within four years, being characterised by markedly higher growth rates than climax species and brief and early fecundities. Climax shrubs had more or less constant rates of mortality over time, though populations in unburnt vegetation tended to have slightly higher rates of mortality than young populations. Densities of seedling populations were very high, but mortality rates were extremely low. In summary, it may be said that the fynbos communi ties studied here are very stable under a given fire regime. Recovery is rather rapid, being apparently achieved within 10 yr. Not much change occurs in older vegetation, but there was a gradual attrition of populations of dominant shrubs, without recruitment, with rare exceptions. Summer droughts in these montane environments are evidently not sufficiently marked for water deficits to play a primary role in succession, so that fire has no effect on plant water relations. Nutrient responses are relatively weak, and masked in the plants by the low rates of metabolism in climax species. Succession after fire is distinguished by the recovery of pre-fire communities, and subsequent inhibition of recruitment. This inhibition is probably through the effects of canopies on microclimate, although the interactions between especially plants and animals have been implicated in succession in other studies. / AC2017

M.Sc. / Please refer to full text to view abstract

The trophodynamic implications of reduced freshwater inflow on the zooplankton of eastern Cape estuaries was investigated by a comparison of the community composition and standing stocks, grazing rates and food resources of zooplankton in two systems, the Kariega and the Great Fish estuaries, which are subject to contrasting freshwater inflow. The climate of South Africa is semi-arid, and the low rainfall, coupled with high evaporative loss, result in the region having one of the lowest conversions of rainfall to run-off in the world. In addition, many of the major rivers are extensively impounded, consequently, estuaries often experience prolonged periods of zero or reduced freshwater inflow. The amount of freshwater available for estuarine management in the future is expected to decline as the demand for domestic, agricultural and industrial use increases. The influence of climate, tidal amplitude and wave action are essentially constant, consequently, the individual characteristics of an estuary are determined largely by the indirect influences of catchment size and regulation. Estuaries along the eastern Cape coast range from negative hypersaline systems, to positive systems in which a salinity gradient is well established. The Kariega estuary is a homogeneous marine estuary as a result of minimal freshwater inflow, whereas the Great Fish estuary receives sustained freshwater inflow and is partially-stratified. The quality and quantity of particulate food resources for suspension-feeders depended to a large extent on the allochthonous import of material associated with freshwater inflow. Nutrients, rather than light penetration of the watercolumn are the major factor limiting phytoplankton standing stocks. In the Kariega estuary, phytoplankton standing stocks were low (up to 1.0 μg l⁻¹) and the estuary is classified as oligotrophic. Correlation analyses indicated that lower quality detritus, originating from fringing macrophytes, may contribute substantially to suspended particulate organic material. Phytoplankton food resources were considerably higher in the Great Fish estuary (up to 21.8 μg ⁻¹) which is classified as mesotrophic/eutrophic, and correlation analyses indicated that phytoplankton comprised the dominant fraction of the particulate organic material. Although this suggested that the organic material was of a higher quality, the seston was "masked" by a higher suspended inorganic load. Consequently, the organic fraction comprised between 13 and 22% of total particulate material in the Great Fish estuary, compared with between 20 and 39% in the Kariega estuary. Food resources demonstrated a fairly uniform distribution along the length of the Kariega estuary and exhibited a slight seasonal increase during warmer months. By contrast, the point source influence of freshwater inflow resulted in a spatial gradient of food resources in the Great Fish estuary with higher values recorded towards the upper reaches. There was evidence that higher concentrations of particulate material in the upper reaches are also a consequence of hydrodynamic trapping. There was no marked seasonal pattern in the availability of food resources which were generally elevated in response to sporadic pulses of freshwater inflow. Tidal currents were responsible for elevating suspended food resources by re-suspension of material from the sediments. This effect was probably of greater importance in the Kariega estuary where food resources were generally limiting. In the Kariega estuary, the zooplankton community was dominated by calanoid copepods of the genera Acartia and Pseudodiaptomus. However, in the Great Fish estuary, the community was dominated by the mysid Mesopodopsis siabberi, and the calanoid P. hessei. Community biomass generally reflected the trophic status of the estuary. The mean seasonal biomass recorded in the Kariega estuary was 38 mg m⁻³ compared with 1597 mg m⁻³ in the Great Fish estuary. Greater spatial variability in community biomass was evident in the Great Fish estuary, partly in response to the food resource gradient, but also due to the inability of the mysid shrimps, which dominated the community biomass, to penetrate the lower salinity water of the upper reaches. Zooplankton in the Kariega estuary demonstrated a seasonal pattern of abundance whereas in the Great Fish estuary, community biomass was elevated in response to sporadic pulses of freshwater inflow. Grazing rates, measured in situ using a modified Gliwicz-Haney chamber, indicated that the zooplankton communities were capable of "clearing" substantial proportions of the watercolumn at certain times of the year. The pattern of grazing pressure over a diel cycle was examined in relation to the diel vertical migration movements of the zooplankton. Higher nighttime grazing rates were generally associated with the greater abundance of zooplankton resulting from the movement of zooplankton into the watercolumn after dusk, and their return to the sediments at dawn. Seasonal estimates of diel grazing pressure, extrapolated from daytime and nighttime feeding rates, indicated that the zooplankton "cleared" up to 40% of the watercolumn in a day in the Kariega estuary, and up to 120% d⁻¹ in the Great Fish estuary. However, values of around 25% d⁻¹ in the Kariega estuary, and 50 to 80% d⁻¹ in the Great Fish estuary, were not uncommon. Multiple regression analyses were used in an attempt to explain the influence of environmental factors on the variation in in situ grazing rates. These attempts were largely unsuccessful and the possible reasons, as well as recommended improvements to the methods used, are discussed. Seston concentration in the estuaries was highly variable as a result of the effects of tidal re-suspension and freshwater inflow. Consequently, further laboratory-based experiments were carried out to examine the influence of seston concentration on the filtration rates of the dominant calanoid copepods. Results indicated that some of the unexplained variability in the community filtration rates may be attributed to differences in species-specific response to changes in seston concentration.

The effects of seawater inflows and macrophyte beds on community metabolism and phosphorus dynamics in the seasonally-closed Swartvlei estuary were investigated. Metabolic rates were determined by diurnal oxygen curve analysis. Gross primary production ranged from 0,7 to 14,9 g 0₂ m⁻² day⁻¹, and respiration from 0,9 to 25,2 g 0₂ m⁻² day⁻¹. The highest production rates were recorded inside the dense beds of Zostera capens is Setchell (x- = 7,8 g 0₂ m⁻² day⁻¹ ). Metabolism was positively correlated to submersed macrophyte cover, and decreased upstream of the mouth region. No seasonal variation in gross production could be detected, and mouth closure had no detectable effect on metabolic rates. Growth of the green alga Enteromorpha sp. in winter, and the effect of decay of this alga on dissolved oxygen, total inorganic carbon and total dissolved phosphorus in the water column is discussed. High day-time respiration values, measured in a darkened polythene enclosure, indicate that photorespiration occurs in Zostera. Apparently the annual amount of oxygen respired throughout the estuary exceeded that produced by 2 7100 tonnes, i.e. the estuary was heterotrophic. This is discussed in relation to the effectiveness of the diurnal curve method as a way of measuring metabolic rates, and to the possibility of organic matter input via river inflows. There is a net gain of up to 17,4 kg of particulate phosphorus over mean tides. Diurnal phosphorus studies indicate that Zostera releases phosphate into the water column in the light. Enteromorpha takes up phosphate in the morning, which it appears to release over the rest of the day. The cycling of phosphorus between seawater, sediments, macrophytes and the water column is discussed.

From Introduction: Energy passes through an ecosystem via a multiplicity of interconnected routes, which can be broadly categorised into trophic and detrital pathways. The "metabolic activity" of most lakes will be governed predcminantly at the base of these two routes; namely, the primary producer and decanposer levels, respectively (Wetzel and Allen, 1972). The importance of the littoral primary producers (especially the aquatic macrophytes) in the functioning of the Swartvlei ecosystem has been emphasised in a comprehensive report by Howard-Williams and Allanson (1978) dealing with the lake system fran 1975 to 1978. They noted that although the littoral shelf (<2m below low water level) occupies only 43% of the lake's surface area it contributed 64% of the total annual primary production during the period investigated. This was largely due to the dense Potamogeton pectinatus stands which alone accounted for 52% of the total carbon input into the lake by plants. The fact that the production/biomass ratio for P. pectinatus was only 1,2:1 suggested that its importance as a primary producer in Swartvlei was largely due to its high bianass. (Biomass, or standing stock, is used in this report as defined by Waters (1977); namely, "the amount present at a point in time, expressed best as quantity per spatial unit".)

M.Sc. (Zoology) / Please refer to full text to view abstract

Investigations were carried out on aspects of the biology of the vetigastropod Turbo sarmaticus. Studies included: 1) the distribution and standing stock of this animal at four sites along the coast of the Eastern Cape Province of South Africa; 2) the growth rate of animals on a wave-cut platform; 3) the reproductive cycle of an intertidal population; and 4) aspects of the feeding biology examining the ability of this mollusc to consume and digest six macroalgae, the influence of algal diet on growth rate and reproductive fitness and the polysaccharolytic activity of the digestive enzymes. On eastern Cape shores, T. sarmaticus had a size related distribution, with smaller animals being found towards the upper mid-shore and larger animals being found in a downshore direction. The mean shore densities of T. sarmaticus at three sites where exploitation of animals was minimal, were very similar (1.2 - 1.7 individuals/m²). The largest animals (up to 110 mm shell length) were found on an offshore island. This may have been a result of animals not being exploited, as well as a possible increase in primary productivity and food availability. The lowest density (0.2 individuals/m²) and animal size (<70 mm shell length) was recorded at a site (Kelly's beach - Port Alfred) where exploitation was more intense. It is probable that intense overexploitation was threatening the populations at this site. The growth rate of T. sarmaticus was determined by means of the von Bertalanffy growth model and expressed by the equation L[subscript]t = 81.07(l-e⁻°·⁵⁴⁴[superscript](t)). The initial growth rate of T. sarmaticus (up to ≈ 80 mm shell length) was similar on shores with different geomorphologies (i.e. boulder shores and wave-cut platforms). Growth rates of individuals were variable, which means that individuals within a population reached exploitable size (3 - 6 years old) and sexual maturity (1.5 - 2 years old) at different ages. Seasonality of reproduction of T. sarmaticus was determined using gonad index, egg diameters and spermatozoa content within the gonad. Turbo sarmaticus was dioecious and had a sex ratio in favour of males (1.2: 1). Animals attained sexual maturity at a size of about 52.5 mm shell length. There was little variation in the reproductive cycle over time with gametogenesis occurring from March/April until August/September, whilst maturity (Gonad Index = 15%) was maintained until the spawning event from December to March. After spawning the gonad regressed. Field and laboratory observations of the feeding biology of T. sarmaticus confirmed that this mollusc was a generalist grazer capable of consuming and digesting algae from the Rhodophyta, Chlorophyta and Phaeophyta. The consumption rates (juveniles: 1.45 - 9.50% body weight/day, adults: 1.06 - 6.08%) and digestibility (9 - 75% apparent dry matter) of six macroalgae was found to vary. For most algae, juvenile T. sarmaticus had higher consumption rates (1.6 - 2.8 times higher) and digestibility values (12 - 24% higher) than adults. It is suggested that consumption rates were dependent on the digestibility of the algae. In addition, it is suggested that the consumption rates of the different algae were not related to the nutritional content, but rather the energetic content of the algae. In both juvenile and adult animals, temperature had a positive influence on consumption rates, resulting in an increase at higher temperatures. However, in both juvenile and adult T. sarmaticus, algal digestibility was not affected by temperature. Finally, it was proposed that Viva rigida, Codium extricatum, Ecklonia radiata and Gelidium pristoides would provide the best nutritional value for growth and reproductive fitness in T. sarmaticus, whilst Jyengaria stellata and Corallina spp. would provide the poorest. Experiments on the effects of four algal diets on the biology of T. sarmaticus showed that the best growth rate (up to 13.8 mm shell length increase per annum), reproductive fitness (Gonad Index up to 33%) and energy levels (up to 4.76% glycogen in the foot) were achieved when T. sarmaticus was fed G. pristoides, U. rigid a or a mixed diet. Turbo sarmaticus fed Corallina spp. showed reduced growth (2.4 mm shell length increase), reproductive fitness (Gonad Index up to 4.4%) and energy levels (up to 3.42% glycogen in the foot). A study of the polysaccharolytic enzyme activity of T. sarmaticus indicated that this mollusc possesses enzymes that can, at least partially, digest most of the storage and structural polysaccharides found in the Chlorophyta, Rhodophyta and Phaeophyta. This further supported the findings that T. sarmaticus was a generalist grazer. Two levels of activity were detected: 1) high levels of enzyme activity (up to 328.2 Ilglmglmllhr)occurred on the storage polysaccharides that occur in the Rhodophyta and Chlorophyta, and 2) lower levels of activity were detected on the storage polysaccharides (up to 44.8 μg/mg/ml/hr) of the Phaeophyta and on all the structural polysaccharides tested (<45.5 μg/mg/ml/hr). It was suggested that T. sarmaticus did not rely heavily on structural carbohydrates as a source of carbon. Finally, the results of this study were discussed in relation to the future management of T. sarmaticus stocks, the possible role of this macro algal grazer in the intertidal zone and the effects of over-exploitation of this animal. The potential aquaculture of this mollusc was also addressed briefly.

The mandatory and voluntary carbon markets have both developed around the increasing trade of carbon offsets. In order to add legitimacy to an otherwise intangible commodity there has been a rise in the development of third party carbon certification standards, particularly in the voluntary market. These standards aim to provide independent, third party certification to projects that are developed specifically to generate and sell carbon offsets. South Africa has the opportunity to engage with these markets, but current participation in and certification of projects is sluggish. These projects have not taken off mainly due to the high transaction costs and lag times surrounding the current certification of projects', complex baseline methodologies, accounting uncertainty and the often bureaucratic systems surrounding the current voluntary carbon certification standards and methodologies. In order to overcome these pitfalls this project aimed to address these challenges through initiating the development of a preliminary South African voluntary carbon standard. This was done by: a) undertaking a critical assessment of the development of current carbon markets, with a particular focus on voluntary markets and third party certification, b) critically analysing the current voluntary carbon certification standards for best practices, pitfalls and weaknesses. To provide a better understanding of the historical development of voluntary certification standards, various established certification standards were evaluated, including the Forestry Stewardship Council (FSC), Marine Stewardship Council (MSC) and ISO 14000 standards. This analysis focussed on the challenges they faced in acceptance and in particular how they have managed to operationalize sustainable development within the certification process. In order to explicitly ensure the incorporation of a sustainable development assessment of projects under the proposed standard, an expert workshop was held with 14 experts from a wide variety of disciplines. These experts identified the crucial sustainable development challenges facing South Africa. They identified 12 sub-themes and 44 indicators that could be used to measure and incorporate sustainable development indicators into the certification process. These were then further developed through using the 'Input – Output – Outcome – Impact' framework model which allowed the indicators to be organised and understood and thus practical. The analysis of the voluntary certification standards and the development of the sustainable development indicator framework were ultimately incorporated into the development of the proposed South African voluntary carbon standard. The key approach to this standard is the incorporation and focus of the proposed standard to ensuring the generation of net SD benefits and placing them at the same level as carbon within the project design and development, validation and verification process. The full and effective integration of these has been missing within current fully fledged voluntary carbon standards, as they often rely on a mixture of project design standards to achieve this. Offering the inclusion of all components into one standard, specifically designed for South Africa, will not only assure SD credentials but also increase transparency and understanding, and reduce costs. This thesis allowed for the development of innovative new ideas and process focussed specifically at including and mainstreaming South Africa’s developmental challenges into the certification process. This is the hoped that the standard will effectively certify South African based landscape restoration projects, but also decrease costs and increase efficiencies in order to encourage the development of these projects. This preliminary standard not only aims to incorporate and address all the issues identified but also has the end goal of acting as the basis for future debate and development surrounding a potential South African voluntary carbon certification standard.

D.Phil. / The results of this investigation suggested that fish hatcheries may cause a deterioration in water quality of thereceiving stream. With regards trout hatcheries, two sources of pollution were distinguished. These were tenned normal or continuous hatchery effluent and secondly dam cleaning effluent. The former effluent was of little pollution consequence. The bacterial analyses indicated that faecal bacterial concentrations were largely determined by the concentration of these bacteria already present in the influent water. However, hatcheries can result in an increase in faecal bacterial concentrations downstream. Biological growth potential investigations suggested that both normal effluents and dam cleaning effluents were only slightly enriched in comparison with the influent conditions. Metabolizable nutrient addition to the effluent water was considered to be of little importance. Although the hatchery effluent from trout hatcheries may result in marked disturbances in the aquatic macro-invertebrate fauna, the organism community structures indicated that severe organic pollution was not occurring. The communities of the macro-invertebrates had become almost normalized within 300 m of the hatchery outfalls'. The water chemistry analyses confirmed the biological findings. The suspended material from both trout and carp hatcheries was the most important contributor to pollution. Recommendations included the use of the existing chemical and physical standards of the Water Act of 1956, however that these should be supplement~d by other biological parameters. A distance of 1 km should exist between a trout hatchery and a farm boundary unless effluent treatment is undertaken. Carp dam effluent may have to be treated depending on the water quality of the receiving stream.

M.Ed. / The purpose of this research was to examine misconceptions held by pupils about Ecology and make suggestions on how these misconceptions could be eliminated. Although there has been much published research identifying misconceptions in the understanding of biological concepts by pupils, relatively little has appeared locally to indicate how misconceptions in this field might best be eliminated, changed or minimized. As part of this research, Biology pupils were given a test to complete. The main aim of the test was to identify misconceptions that the pupils held after classroom instruction. The pupils were also given a questionnaire to complete. The questionnaire was designed to assess pupils' attitude towards Biology in general, Ecology and fieldwork in particular. The pupils' knowledge of Ecology following instruction was further assessed by interviews. Teachers were also given questionnaires to complete. The main aim of the teachers' questionnaire was to determine if fieldwork could play any significant role in the teaching of Biology. Furthermore, the teachers' questionnaires also explored some of the problems or limiting factors experienced by teachers when they undertake fieldwork. This study revealed that pupils held misconceptions about energy flow in the ecosystem, basic ecological terms/concepts, nutrient cycling, the food web, etc. The research also suggested that fieldwork helps pupils to understand certain other sections of the Biology syllabus better. It is recommended, amongst others, that experienced Biology teachers, authors of Biology textbooks, lecturers at in-service training centres, and subject advisors should do all they possibly can to assist their colleagues and pupils to avoid misconceptions about Ecology.