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. 2016. / Solanum mauritianum Scopoli (Solanaceae) is a perennial tree or shrub native to South America, which has become a prominent and widespread invader in numerous sub-tropical countries around the world. In South Africa, S. mauritianum is listed as one of the country’s worst ecological weeds, having been targeted for biological control efforts since 1984. Despite some constraints, biocontrol efforts have seen the successful release of two promising biocontrol agents. The first of these biocontrol agents, released against S. mauritianum, was the sap-sucking lace bug, Gargaphia decoris Drake (Hemiptera: Tingidae). Sap-feeding by G. decoris metabolically impaired the leaves, resulting in a reduction to their photosynthesis, with a greater effect on plants growing in full-sun compared to plants growing in the shade. This difference was attributed to higher leaf temperatures experienced in the sun. Herbivory reduced transpiration rates by more than 50%, resulting in a reduction in evaporative cooling of the leaf. The increased physiological damage experienced by full-sun plants may be a combination of stresses, particularly the direct effect of chlorophyll removal via herbivory and the indirect effect of accumulated heat–light stress. The flowerbud-feeding weevil, Anthonomus santacruzi Hustache (Coleoptera: Curculionidae), was released in 2008 as a biological control agent against S. mauritianum. The hypothesis that climate, particularly low temperature and low relative humidity, restricts the survival and establishment of A. santacruzi in South Africa was tested. Thermal assessments on A. santacruzi adults calculated the CTmin and LT50 as 4.1 ± 0.2 °C (n = 20) and 4.2 ± 0.3 °C (n = 90) respectively. The LH50 of A. santacruzi adults was calculated as 46.9%. The establishment of A. santacruzi at only the warm and humid release sites in South Africa advocates for the consideration of low temperature and low humidity as factors impeding the agents’ establishment and spread, particularly on the cooler and drier Highveld. Furthermore, the impact of A. santacruzi’s florivory on the reproductive output of S. mauritianum, as well as the potential of the agent to act as an indirect pollinator was assessed. Overall direct floral damage caused by A. santacruzi was trivial, with only ~5% of the anther and ~2% of the petal area being removed. However, the consequent effects of A. santacruzi were considerably more damaging, with 25% and 66% reductions in flowering and fruiting respectively. Additionally, fruits produced from inflorescences exposed to A. santacruzi were smaller in size, with fewer, less viable seeds. The feeding and presence of A. santacruzi also maintains the potential for indirect effects on the pollination of S. mauritianum. This suggests that in areas with well-established A. santacruzi populations, the weevils may simultaneously facilitate the self-pollination and potential inbreeding of S. mauritianum. Keywords: Agent impacts and effects; biological control; Bugweed; climatic unsuitability; ecophysiology; indirect effects; post-release evaluation. / LG2017

Thesis (MSc)--University of Stellenbosch, 2005. / ENGLISH ABSTRACT: This project consists of two botanical investigations in an agricultural area of the Western Cape Province. A farm known as De Rust, in the Elgin Valley, was used to sample the geographic location, density, height and life stage of six prominent invasive plant species in various land-use categories. In the first investigation, the density, height and age structures of the six invasive species populations were analyzed. The density distribution of the six species was also displayed cartographically. Species were then ranked according to the potential threat that they pose to the conservation of the remaining natural areas on the farm. Results indicated that Acacia mearnsii and Acacia saligna are the major invaders at De Rust and that Hakea sericea can be considered as an emerging invader. The second investigation explores the statistical relationship between the various land-use categories and density, height and age of the six prominent invaders identified in the first investigation. The loglikelihood ratio analysis of observed frequencies resulted in statistically significant (P<0.01; P-values range between 1.35 x 10-3 and 2.7 x 10-224) relationships between certain land-use types and certain invasive species. A conclusion was reached that it could be useful to include land-use categories in simulation models of invasive plant species distribution and spread. / AFRIKAANSE OPSOMMING: Hierdie projek behels twee botaniese ondersoeke in ‘n landbou gebied van die Weskaap. Die plaas bekend as De Rust, in die Elgin Vallei, was gebruik vir die versameling van data te doen met die geografiese ligging, plant digtheid, lengte en lewens stadium van ses prominente indringer plant spesies in verskeie landgebruik kategorieë. Die digtheid, lengte en ouderdomstruktuur van ses indringerspesies was in die eerste ondersoek geanaliseer. Die verspreiding van digtheid was ook in kaarte uitgelê. Spesies was daarna volgens hulle potentiële dreiging teen die bewaring van oorblywende natuurlike dele van die plaas in ‘n rangorde geplaas. Resiltate dui aan dat Acacia mearnsii en Acacia saligna die belangrikste indringer plante op De Rust is en dat Hakea sericea as ‘n opkomende indringer beskou kan word. Die tweede ondersoek kyk na die verhouding tussen verskeie grondgebruik kategorië en die digtheid, lengte en ouderdom van die ses prominente indringer spesies wat in die eerste ondersoek identifiseër is. ‘n Log tipe ratios ontleding van bewaarde frekwensies het ‘n statisties belangrike uitkoms gehad (P<0.01; P-waardes tussen 1.35 x 10-3 en 2.7 x 10-224) vir die verhoudings tussen sekere grondgebruik tipes en sekere indringer spesies. Die gevolgtrekking was dat dit handig mag wees om grondgebruik kategorieë in simulasies van indringer plant verspreiding te gebruik.

Water hyacinth, Eichhornia crassipes (Martius) Solms-Laubach (Pontederiaceae), a free floating aquatic plant was discovered by C. von Martius in 1823 in Brazil. It is believed to have been introduced into South Africa, as an ornamental plant, in 1908 to the Cape Province and Natal. Since its introduction, water hyacinth has spread throughout South Africa to the detriment of all aquatic systems that it has been introduced to directly or indirectly. The weed was first positively identified on the Nseleni and Mposa rivers on the Nseleni Nature Reserve which is a protected area near Richards Bay in KwaZulu- Natal in 1982 and formed a 100% cover of the river by 1983. An integrated management plan was implemented in 1995 and resulted in a reduction of the weed from a 100% cover to less than 20% cover in 5 years. The keys to success of the water hyacinth integrated management plan, presented here, were finding the source of the weed, mapping the extent of the water hyacinth infestation, identifying sources of nutrient pollution, appointing a champion to drive the programme, dividing the river into management units, consultation with interested and affected parties, judicious use of herbicides and biological control and a commitment to follow-up. This study further showed that water hyacinth on the Nseleni and Mposa river systems had a negative impact on the biodiversity of the protected area and the control of water hyacinth resulted in the recovery of the benthic invertebrate, amphibian, reptile, fish and avian fauna. The implementation of this integrated management plan was very cost-effective and serves as a model approach to the control of water hyacinth in both South Africa and the rest of the world.

Aquatic weeds transform and degrade the ecosystems which they invade, impacting various aspects of their surroundings ranging from the community level to disrupting important processes affecting ecosystem services. All of the major aquatic weeds of South Africa are found in the Cape Town Metropolitan Area. Landowners, whether private or public, are legally obliged to manage the listed invasive species through applying environmentally acceptable methodologies. This thesis provides an overview of the strategic management options, prevention, early detection, rapid response and eradication of new invasions, and containment and control species of established species. It discusses the different control methods available for managing aquatic weeds, namely mechanical, manual, chemical and biological, and the integration of different methods to improve their effectiveness. Although various studies have shown that biological control is the most cost–effective, environmentally-friendly and sustainable method, it is not yet fully integrated into weed management programmes in South Africa. In addition, the successes achieved in other parts of the world with the control of water hyacinth through biological control have not been repeated in the urban environment, despite the fact that South Africa has the highest number of biological control agents available for the weed. Urbanisation puts pressure on the natural environment and ecosystem functioning. Nutrient-enriched waters support aquatic weed growth and pose a challenge to the management thereof, in particular with regard to integrating biological control into management programmes. The aims of this study were to determine the reasons for the lack of integration of biological control into weed management programmes in South Africa, to determine the feasibility of integrating biological control in aquatic weed management programmes in a complex urban environmental and socio-political landscape by means of three case studies in the Cape Town Metropolitan Area, which showed that biological control is feasible in urban environments and should be considered. Two surveys were conducted to determine the reasons for the lack of integration of biological control into weed management programmes. The surveys showed that there is a gap between research and implementation as a result of poor communication, non-supporting institutional arrangements and a lack of appropriate capacity and skills at the implementation level. Recommendations were offered to address these issues.

An autecological study of the heterosporous fern Azolla filiculoides Lamarck and its endosymbiotic blue-green alga Anabaena azollae Strasburger, based on a combination of field and laboratory studies, is presented. The taxonomy, morphology and anatomy of the fern-alga association were studied as well as nutritional and physiological aspects of the symbiosis. These studies have defined the habitat and nutritional requirements of the fern and have provided new insights into its reproductive biology, nitrogen metabolism and the nature of the association between the fern and alga. In the catchment area of the Hendrik Verwoerd Dam the availability of suitably sheltered habitat limits the distribution of A. filiculoides while the availability of nutrients, in particular calcium, phosphorus and iron, limits the growth of the fern. The multilayered mats formed by A. filiculoides are essential for spore production, cause dramatic changes in the hydrochemistry of the underlying waters and confer a great competitive advantage on the plant. Methods for the isolation of the fern and algal components of the symbiosis have been developed but recombination of the individual organisms to reform the symbiosis was unsuccessful. The development of the fern is closely linked to that of the alga and the association is maintained throughout the life cycle of the fern. Because of its specific habitat and nutritional requirements, A. filiculoides is unlikely to colonize the open waters of the Hendrik Verwoerd Dam.

MSc (Botanty) / Department of Botany / Background: Problem plants and alien invasive weed species are part of today‟s ecological transformation. Ethnobotanical literature on ecological prominence and ethical values of problem alien weeds is scant and should be documented to avoid loss of valuable species. Aim: The study was aimed at documenting problem weeds and alien invasive plant species considered to be beneficial in deep rural communities. Materials and method: Purposeful triangulation research methods were used to ensure the proper gathering of both qualitative and quantitative data sets. Results and conclusion: Some problem weeds and alien invasive plant species are being utilized by local people to maintain their livelihood. A total of 78 plant species from 33 families were recorded. These plant species belonged to diverse growth habits, namely: climbers, herbaceous, succulents, shrubs, trees as well as creepers. Recorded problem weeds and alien invasive plant species were contributing towards ecological, medicinal and social values. These included being utilized as a source of food (14), fruits (4), medicinal (31), ornamentals (7), firewood (3), social (2) and multiple purposes (17). The main utilized families were Asteraceae (14.1%) and Solanaceae (14.1%). These families were mostly utilized for the purpose of medicinal and food. Plant species that were widely used are as follows: Achyrathes aspera L., Agave sisalana L., Amaranthus spinosus L., Anredera cordifolia L., Canna indica L., Centella asiatica L., Chenopodium album L., Chenopodium ambrosioides L. and Chromolaena odorata (L.) R. M. King and H. Rob. The dominant category for problem weeds and alien invasive plant species was 1b (44%) followed by weeds (35%), category 2 (9%), invaders (5%), category 3 (4%) and 1% for both category 1, 1a as well as X3. The results revealed that the legislative listing of plant species as problem weeds and alien invasive species was based on single knowledge basis system, rather than on multi-dimensional knowledge systems. Therefore, this study recommended that for judgment to be considered in decision-making, it should be based on species-specificity as well as multi-dimensional-knowledge basis. The reconciliation of inherent grapples among scientific and indigenous knowledge systems could possibly be underpinned by equal legislative consideration for the aforementioned knowledge systems especially in the establishment and implementation of environmental regulations. This could also aid with the provision to support transformation in South Africa and worldwide.

The invasion of ecosystems by alien species is a growing threat to the delivery of ecosystem services. This study explored the spatial distribution of water hyacinth in the Benoni Lakes and made analysis of its impact on water availability and usability in order to understand the evolution and its propagation rates. The study used satellite imagery for lake level modelling using Remote Sensing and Geographical Information Systems for calculations of area covered by weeds in each lake from 2002 to 2012.The modelling approach illustrates the potential usefulness in projecting invasive plants under climate change and enabled the quantification of long term changes in aquatic weeds. The results showed that aquatic infestations in lakes may be used as powerful predictors of correlations between plant abundance and climate change. The study therefore informs decision makers to identify areas where invasion is likely to occur and increase surveillance for early invaders. / Environmental Sciences / M. Sc. (Environmental Management)

Water hyacinth (Eichhornia crassipes) is an aquatic macrophyte originating from the Amazon basin. Due to its beautiful appearance it has been introduced into numerous countries across the world as an ornamental pond plant. It was introduced into South Africa in the early 1900s and has since reached pest proportions in many of the country’s fresh water bodies, causing significant economic and ecological losses. It is now considered to be the worst aquatic weed in South Africa. Efforts to control the spread of the weed began in the early 1970s and there have been some successes. Biological control has been used widely as an alternative to mechanical and chemical controls because it is cost-effective, self-sustaining and environmentally friendly. To date, six biological control agents have been introduced onto water hyacinth in South Africa. However, due to factors such as cold winter temperatures and interference from chemical control, the agent populations are occasionally knocked-down and thus the impact of biological control on the weed population is variable. In addition, many South African water systems are highly eutrophic, and in these systems the plant growth may be accelerated to such an extent that the negative impact of the agents’ herbivory is mitigated. One of the agents established on the weed is the galumnid mite Orthogalumna terebrantis, which originates from Uruguay. In South Africa, the mite was initially discovered on two water hyacinth infestations in the Mpumalanga Province in 1989 and it is now established at 17 sites across the country. Many biological control researchers believe that the mite is a good biological control agent but, prior to this thesis, little quantitative data existed to confirm the belief. Thus, this thesis is a post-release evaluation of O. terebrantis in which various aspects of the mite-plant relationship were investigated to determine the efficacy of the mite and thus better understand the role of the mite in the biological control programme of water hyacinth in South Africa. From laboratory experiments, in which mite densities were lower than densities occurring in the field, it was found that water hyacinth growth is largely unaffected by mite herbivory, except possibly at very high mite densities. When grown in high nutrient conditions the growth of the plant is so great that any affect the mite has is nullified. Plant growth is thus more affected by nutrients than by mite herbivory. However, mite feeding was also influenced by water nutrient levels and mite herbivory was greatest on plants grown in high nutrient conditions. The presence of the mite had a positive effect on the performance of the mirid Eccritotarsus catarinensis, such that the interactions of the two agents together had a greater negative impact on the plant’s growth than the individual agents had alone. Furthermore, water hyacinth physiological parameters, such as the plant’s photosynthetic ability, were negatively impacted by the mite, even at the very low mite densities used in the study. Plant growth rate is dependent on photosynthetic ability i.e. the rate of photosynthesis, and thus a decrease in the plant’s photosynthetic ability will eventually be translated into decreased plant growth rates which would ultimately result in the overall reduction of water hyacinth populations. In addition, temperature tolerance studies showed that the mite was tolerant of low temperatures. The mite already occurs at some of the coldest sites in South Africa. Therefore, the mite should be able to establish at all of the water hyacinth infestations in the country, but because it is a poor disperser it is unlikely to establish at new sites without human intervention. It is suggested that the mite be used as an additional biological control agent at sites where it does not yet occur, specifically at cold sites where some of the other, less cold-tolerant, agents have failed to establish. Finally, conditions of where, how many and how often the mite should be distributed to water hyacinth infestation in South Africa are discussed.

Potentially damaging submerged invasive freshwater macrophytes have been identified in South African freshwater systems, but have received less attention than their floating counterparts. To ascertain the changes and effects that these species may have on macrophyte ecology, an understanding of the drivers of macrophyte assemblages is essential. The aims of this thesis were to investigate select abiotic and biotic factors driving introduction, establishment and spread of submerged macrophytes in South Africa. Surveys on the status of submerged plant species in South Africa were conducted to find out the distribution and diversity of the species present, imported to, and traded in South Africa. Numerous submerged indigenous and invasive macrophyte locality records were collected during field surveys, of which many were first time records. Pet stores and aquarist trading activities were identified as potential vectors for the spread of submerged macrophytes through online surveys and personal interviews. These results highlighted the potential these species have for continuing to enter, and spread within South African water bodies. Maximum Entropy (MAXENT) is a general-purpose method used to predict or infer distributions from incomplete information, and was used here to predict areas suitable for the establishment of five of these invasive macrophytes. Many systems throughout South Africa, particularly those in the subtropical coastal regions, were found to be climatically suitable for the establishment of Elodea canadensis Michx., Egeria densa Planch., Hydrilla verticillata (L.f.) Royle (all Hydrocharitaceae), Myriophyllum spicatum L. (Haloragaceae), and Cabomba caroliniana Gray (Cabombaceae). Despite the high probability of invasion, facilitated by vectors and suitable climate, South Africa’s rich indigenous submerged aquatic flora may be preventing the establishment of these submerged invasive species. Studies on the competitive interactions between a common indigenous submerged macrophytes, Lagarosiphon major (Roxb.) (Hydrocharitaceae) and M.spicatum, an invasive native to Eurasia, were conducted to ascertain which conditions influence competitive superiority. High sediment nutrient conditions significantly increased the growth rate and competitive ability of both species, while clay sediments significantly increased the competitive ability of L. major over M. spicatum, but sandy sediments improved the competitive ability of M. spicatum. These results highlighted the dynamic changes in competition between submerged species driven by abiotic factors, but did not take into consideration the effect that herbivory, a biotic factor, could have on competition between the two species. The effect of herbivory by phytophagous insects of submerged plant species has been regarded as negligible. To find out what this effect is, multiple field surveys were undertaken throughout South Africa to find natural enemies of indigenous Lagarosiphon species with the aim of identifying such species, and quantifying their influence on plant growth dynamics. Several new phytophagous species were recorded for the first time. An ephydrid fly, Hydrellia lagarosiphon Deeming (Diptera: Ephydridae) was ascertained to be the most ubiquitous and abundant species associated with L. major in South Africa. The influence of herbivory by this fly on the competitive ability of L. major in the presence of M. spicatum was investigated using an inverse linear model, which showed that herbivory by H. lagarosiphon reduced the competitive ability of L. major by approximately five times in favour of M. spicatum. This study served to highlight the importance of herbivory as a driver of submerged aquatic plant dynamics. Current ecological theory emphasises the importance of investigating beyond plant-herbivore interactions, by including multitrophic interactions in community dynamics. Therefore, the potential of parasitism by a parasitoid wasp, Chaenusa luteostigma sp. n. Achterberg (Hymenoptera: Braconidae: Alysiinae) on H. lagarosiphon to shift the competitive interactions between the two plant species was also examined. The addition of the parasitoid reduced the effect of herbivory by the fly on L. major by half, thereby shifting the competitive balance in favour of L. major over M. spicatum. This study provides valuable insight into a selection of drivers of submerged macrophyte assemblages of South Africa. It highlights the precarious position of South African freshwater systems with regard to the potential invasion by damaging submerged invasive species. It also provides interesting insights into the effect of competition, herbivory and parasitism on the establishment and spread of species within submerged freshwater systems. Understanding the different influences could assist managers and policy makers to make validated decisions ensuring the integrity of South African freshwater systems.

The contamination of heavy metals in the environment is a looming concern worldwide. Egeria densa (Planch) (Submerged aquatic plant) from two ponds: Site A with co-ordinates (32º 48’22.04”S; 26°48’58.79” E) and Site B with co-ordinates (32°48’33.25”S; 26°48’33.25”S) in Alice (Eastern Cape) was evaluated for its ability to absorb heavy metals, phytochemical constituents, antimicrobial activity and ultra-structure using standard analytic procedures. Cadmium (Cd), copper (Cu), iron (Fe), lead (Pb), manganese (Mn), and zinc (Zn) were measured in water, sediments and plant. The concentrations of these metal elements were determined with use of Inductively Coupled Plasma- Optical Emission Spectrometry (ICP-OES). In sediments, the heavy metals (mg/kg) decreased in the order of their average concentration as follows: Fe (40.320) > Zn (1.259) > Pb (0.564) > Mn (0.186) > Cu (0.037) in Pond 1 whereas in Pond 2 Fe (61.527) > Cd (0.999) > Mn (0.648) > Pb (0.586) > Zn (0.156) > Cu (0.045). The highest concentration of Fe was detected in both sites and Cu being the least. The concentrations of the metals in the plants sample (from Pond 1) were found in order of Mn > Pb > Cu > Fe whereas cadmium and zinc were not detected, while the concentration in Pond 2 decreases in order of Zn > Mn > Pb > Cd > Fe > Cu. In the water samples, concentrations of heavy metals (mg/L) decreased in the order of their average concentrations as follows: Pb (35.36) > Fe (3.07) > Mn (0.238) > Cu (0.104), both cadmium and zinc were below the limit of detection in Pond 1, whereas in Pond 2 the concentrations decreased as follows: Pb (13.033) >Fe (1.69) > Cu (0.270) > Mn (0.248) > Cd (0.004) and Zinc was not detected. Phytochemical analyses of the plant extracts revealed the presence of phenols, flavonoids, proanthocyanidin, flavonols, saponins, alkaloid and tannins in all the extracts (water, acetone and n-hexane). Both acetone and water extracts, showed high concentration of proanthocyanidin, while tannin was the lowest in acetone extract. Antimicrobial evaluation using, Gram positive (Staphylococcus aureus, Bacillus pumilus, Bacillus cereus, Streptococcus pyogenes, Enterococcus faecalis) and Gram negative (Klebsiella pneumonia, Escherichia coli, Pseudomonas aeruginosa, Proteus vulgaris and Serratia marcescens) bacteria showed negative results for all the strain, except Streptococcus pyogenes which was inhibited at MIC of 0.1 mg/ml. Scanning electron microscopy (SEM) of ultra-structure of Egeria densa, showed that certain bacteria attached to the leaf, However more work has to be done on E. densa to verify the mechanism by which it accumulates heavy metals. The study shows that E. densa has a potential of accumulating heavy metals especial Manganese in plant.