This study investigates the economic impact of the ARC PPRI Weeds Research Division. The Division researches appropriate methods of biological control for invasive alien plants (IAPs). These plants pose an increasing threat to environmental integrity and ecosystem service provision impacting on economic potential. Since the work of the Division is considered a public good, a predominantly descriptive approach has been adopted for the valuation process. A combination of quantitative cost analysis and a qualitative study of the impacts of research and invasive alien plants is used to deal with the challenges associated with non-market valuation. The study found that investment into the Weeds Division is a valuable activity that supports the long-term growth potential of the South African economy. The role of a well-functioning environment is highlighted as an essential base for the creation of sustained growth opportunities in any society. It was determined that investment into the Division should be increased into the future to support efficient spending of scarce state funds. Biological control research was found to provide strategic future growth potential, creating opportunities for the development of a competitive advantage in the biotechnology and environmental management sectors. The study adds to the increasing move towards a more holistic view of economic valuation, taking factors other than pure finance and econometrics into consideration. This is an important shift in prevailing economic thought, as a realisation is reached that a single, or even triple, bottom line is an outdated and insufficient decision making basis.

The aims of the project were twofold. The initial objective of the study, based on previous results, was to develop an economically viable methodology for immobilizing yeast cells for the treatment of heavy metal-laden waste water. The non-viable yeast cross-linked by 13% (w/v) formaldehyde/1N HNO₃ exhibited satisfactory mechanical strength and rigidity in a continuous-flow column operation. No apparent disruption of the biomass after repeated use was observed. The cost of immobilizing 1kg dry yeast pellets was estimated at less than US$I. Zn uptake capacity of FA-cross-linked pellets, on batch trials, remained similar to that of raw yeast, reflecting that the immobilizing procedure did not hinder its metal removing capacity. In column studies, cation metals were effectively removed by the yeast pellets from aqueous solution at natural pHs, and then recovered completely by washing the pellets in situ with O.1M HCl. The recovered metals were concentrated in such small volumes that recycling or precipitation of them was facilitated. The metal uptake capacity of the regenerated biomass remained constant in comparison with cycle 1, indicating that reuse of the yeast would be possible. In the case of Cr⁶⁺, a gradual breakthrough curve of Cr in the column profile was noted, with a simultaneous reduction of Cr⁶⁺ to Cr³⁺. However, Cr⁶⁺ in the effluent can be markedly minimised either by accumulation onto the biomass or reduction to its trivalent form. Desorption of bound Cr⁶⁺ with either alkali or salt could not accomplish the regeneration of the biomass. A combination of reduction and desorption with FA/HNO₃ appeared promising in regeneration of the saturated biomass at 4°C. The metal sorption capacities of the yeast pellets, on a batch or a fixed-bed system are relatively lower than that of documented sorbents. Apparently more of the yeast pellets would be required for treating a certain volume of waste effluent, than with other sorbents. Therefore Azolla filiculoides was examined as a suitable sorbent for this purpose. This constitutes the second part of the project. Azolla filiculoides, a naturally-abundant water fern, was screened for its metal sorption and recovering capacities, mechanical stability, flow-permeability and reusability. The azolla biomass appeared to have fulfilled the required mechanical criteria during the repeated sorption-desorption column operations. It is water-insoluble and appears flexible under pressure when rinsed with water. These characters are of crucial importance in a continuous-flow system since a column can be operated at high flow rates without apparent compact of the biomass and pressure loss. Therefore, immobilization of the biomass can be avoided. The sorption isotherm data, obtained from batch removal of Cr⁶⁺, showed that the sorption process was effective, endothermic and highly pH dependent. Considerable amounts of Cr⁶⁺ were accumulated at the optimum pHs of 2-2.5. Column sorption of Cr⁶⁺ at a low flow rate and pH of 2.5 showed optimum performance with a total Cr uptake of 50.4mg/g at 60% saturation of the biomass. Removal of Cr⁶⁺ from an electroplating effluent using an azolla column was deemed reasonably satisfactory, although the uptake declined slightly. Desorption of bound Cr⁶⁺ with various desorbents was incomplete, which resulted in a low regeneration efficiency of about 50%. However, removal and recovery of Cr³⁺ using the azolla column was than that of Cr⁶⁺. Desorption of Cr³⁺ from the spent biomass column was accomplished with the recovery of 80% using O.5N H₂SO₄, The regeneration efficiencies for Cr³⁺ removal were up to 90% and demonstrated that the biomass is reusable. Cation metal uptake capacities of azolla, obtained either from batch or column experiments, are reasonably high in comparison with other sorbents. The uptake of Ni or Zn ions from solution is pH dependent showing the optimum pH of around 6 to 6.5, under the current experimental conditions. The sorption kinetics for cation metals was rapid with about 80% of the bound Ni ions being taken up in the first 10 min. The character of rapid binding is extremely important in a column sorption process, especially on a large scale since it favours an optimum uptake of metals at high flow rates. The Ni or Zn uptakes in column sorption were not markedly affected when the flow rates were increased from 80mllh up to 800ml/h for the 5g biomass used. The cation heavy metals removed from waste effluents were recovered in a concentrated solution of small volume. The desorption of bound Ni and Zn ions from the saturated biomass was accomplished with either O.2N HCl or H₂SO₄ that resulted in recoveries of more than 95%. The metals recovered, in the case of Ni and Zn, are identical to that of plating agents ego nickel sulphate or chloride, so that recycling of the metals is possible. An effluent-free, closed loop of Ni or Zn treatment system was proposed, whereby the Ni or Zn ions can be recycled to the plating bath whilst the purified water is fed back to the rinse tanks. Ca and Mg ions, commonly present in the electroplating effluents, appeared to affect sorption of heavy metals by azolla when metal concentrations were relatively low, presumedly through its competitive binding for the shared sites on surfaces of azolla. The data obtained from column sorption of Ni and Zn follows the BDST model well, enabling the application of the model to predicting design parameters for scale-up of the biosorption column system. It is interesting that the values of metal uptake, expressed in molar quantities, obtained on respective single-metal solutions and the multiple metal system, are similar, implying that the mechanisms involved in the sorption of all metal cations are similar and that the binding sites on surfaces of azolla are probably shared by all cation metals. The surface of the biomass provides sites for metal binding estimated in the range of 0.45-0.57mmol/g, based on the current experiments. The biomass has a surface area of 429 m²/g and water retention of 14.3 ml/g. The functional groups on the surface of azolla were partially identified using chemical modification and metal binding comparison. Among the functional groups examined, carboxyl groups, provided by amino acids and polysaccharides, appeared to play an important role in metal cation binding. The infrared spectra of the samples support this conclusion.

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, March 2017. / Water hyacinth (Eichhornia crassipes) is an example of an alien invasive plant, threatening ecosystems around the world. Taking into consideration all the negative impacts caused by the proliferation of water hyacinth, many control measures have been implemented to manage this weed. These include mechanical control, chemical control and biological control. But in recent years, the focus has shifted towards natural enemies for control (biological control) of water hyacinth because they have many advantages for environmental safety. However, in South Africa, success is limited in most parts of the country. Factors affecting the success of biological control of water hyacinth in South Africa have been attributed in part to weather conditions. Therefore, further studies and other strategies are needed to reinforce the biological control of water hyacinth. This is why this research investigated the influences of temperature on Neochetina eichhorniae larvae feeding and development on water hyacinth, to contribute to the future management of water hyacinth biological control. To measure the effect of temperature on the feeding and development of Neochetina eichhorniae larva, experiments were conducted at four fixed temperatures (15 °C, 20 °C, 25 °C, and 30 °C). Larval feeding is expected to be greater and the development faster at warm temperatures compared with cold temperatures. The results of this study confirmed that, both feeding and larval development increased with an increase in temperature. The highest larval feeding damage was recorded at 30°C and the lowest at 15°C. Similar to the feeding of larvae, warmer temperatures were more favourable to the growth of larvae than cold temperatures; all larvae found at 30°C were in their third instars (with 0.72 mm being the biggest headwidth found) while all larvae at 15°C were in the first instars (with 0.28 mm being the smallest headwidth found). Even if the optimal temperature for larval performance took place at 30 °C (with a high amount of biomass removal), it was at 25 °C that the larvae had a larger effect on the growth of water hyacinth. Larval feeding damage reduced the growth of water hyacinth by 30 % at 25 °C and only affected 24 % of water hyacinth growth at iv 30 °C. Therefore, for the biocontrol of water hyacinth, the ideal temperature to slow the progression of the weed will be recommended at between 25- 30 °C. / LG2017

No description available.

Yellow toadflax, Linaria vulgaris (L.) Mill. (Plantaginaceae), is a non-native invasive plant. Rhinusa pilosa Germar (Coleoptera: Curculionidae) is a proposed biocontrol agent. Gall development by R. pilosa was described using histological methods and compared between plant populations from native and introduced ranges. Key stages of oviposition were isolated histologically to determine their importance in gall induction. Rhinusa pilosa galled and developed on four geographically distinct Canadian populations in a pre-release quarantine study. Low agent densities only negatively affected one population. High densities of R. pilosa reduced potential reproductive output and plant biomass. Conducting detailed investigations into the biology, impact, and development of R. pilosa on populations from invasive and native ranges may help predict the efficacy of R. pilosa in the field if approved for release and.goes beyond current pre-release testing requirements. / ix, 168 leaves : ill. (chiefly col.) ; 29 cm

[Truncated abstract] In the crop growing areas of Western Australia, two economically important weed species, Lolium rigidum Gaud. (annual ryegrass) and Raphanus raphanistrum L. (wild radish), have evolved widespread herbicide resistance to multiple chemistry groups. Consequently, grain growers in the region have adopted an integrated approach to weed management that includes many non herbicide tools, however many more are needed as these weed species become increasingly more difficult to control. This thesis examines, in a series of field trials carried out in the Western Australian crop growing area, the potential for weed seed predation of annual ryegrass and wild radish by naturally occurring granivores as a new weed management tool for grain growers . . . The study discusses the implications of these results with the view to manipulating predation of weed seed through agricultural management practices. Ants were shown to be the dominant seed predator in this environment, especially in the centre of fields. The study has identified that the ant species Melophorus turneri (Forel), Monomorium rothsteini (Forel), Pheidole hartmeyeri (Forel) and Rhytidoponera metallica (Smith) are potential biological control agents for annual ryegrass seeds while P. hartmeyeri was identified as the only species suitable for biological control of wild radish seed pods. Ants were found to be sensitive to disturbance and some to crop residue type and these effects are discussed in relation to seed removal. This study of weed seed predation in agricultural fields is the most complete in this environment and can be used to inform further work in this area. It has identified that naturally occurring granivores can be used as a weed management tool.

Lantana camara (lantana), a thicket-forming shrub, a number of different varieties of which were introduced into South Africa as ornamental plants but which has become a serious invasive weed. Conventional control measures for lantana are expensive and ineffective and it has therefore been targeted for biological control since 1961. To date, eleven biological control agent species have become established on lantana in South Africa. However, most agents persist at low densities and only occasionally impact plant populations. Three species regularly cause significant damage, but only reach sufficiently high numbers by midsummer after populations crash during the winter. Overall, the impact of the biological control programme on the weed is negligible and this has been ascribed to the poor selection of agents for release, the accumulation of native parasitoids, differences in insect preference for different varieties of the weed and variable climatic conditions over the weed’s range. This study suggests that the importance of varietal preferences has been over-estimated. A predictive bioclimatic modelling technique showed that most of the agents established in South Africa have a wide climatic tolerance and that the redistribution and importation of new climatypes of these agents will not improve the level of control. Additional agents are required to improve the biocontrol in the temperate conditions, and also to increase damage in the sub-tropical areas where most of the agents are established and where the weed retains its leaves year round. New candidate agents that possess biological attributes that favour a high intrinsic rate of increase, a high impact per individual and that improve the synchrony between the weed and the agent in climatic conditions that promote the seasonal leaflessness of plants should receive prior consideration. A survey in Jamaica indicated that additional biological control agents are available in the region of origin but that care should be taken to prioritise the most effective agents. The various selection systems currently available in weed biocontrol produce contradictory results in the priority assigned to candidate agents and a new selection system is proposed. The biology and host range of two new candidate natural enemies, the leaf-galling weevil, Coelocephalapion camarae and the leaf-sucking mirid, Falconia intermedia were investigated for the biocontrol of lantana. The studies indicated that these have considerable biocontrol potential, in that the weevil has a wide climatic tolerance and has the potential to survive the host leaflessness typical of temperate conditions, while the mirid has a high intrinsic rate of increase, and the potential for several generations a year. Both agents caused a high level of damage to the leaves, with the weevil galling the vascular tissue in the leaf-petiole and the mirid causing chlorotic speckling of the leaves. During laboratory trials both agents accepted indigenous species in the genus Lippia. However, under multiple choice conditions these agents showed a significant and strong oviposition preference for lantana. A risk assessment and post release field trials indicated that F. intermedia is likely to attack some Lippia species in the presence of lantana, but the levels of damage are predicted to be relatively low. A possible low incidence of damage to indigenous species was considered a justifiable ‘trade-off’ for the potentially marked impact on L. camara. Preference and performance studies on the two candidate agents suggested that most of the South African lantana varieties are suitable host plants. The mirid preferred certain varieties in multiple choice experiments, but this is unlikely to affect its impact under field conditions. Permission for release was accordingly sought for both species. Finally, the challenges facing the biological control programme and the potential for improving the control of L. camara in South Africa are considered.

Water hyacinth, Eichhornia crassipes Mart. Solms-Laubach (Pontederiaceae), a freefloating aquatic macrophyte of Neotropical origin, was introduced into South Africa as an ornamental aquarium plant in the early 1900’s. By the 1970’s it had reached pest proportions in dams and rivers around the country. Due to the sustainability, cost efficiency and low environmental risk associated with biological control, this has been a widely used method in an attempt to reduce infestations to below the threshold where they cause economic and ecological damage. To date, five arthropod and one pathogen biocontrol agents have been introduced for the control of water hyacinth but their impact has been variable. It is believed that their efficacy is hampered by the presence of highly eutrophic systems in South Africa in which plant growth is prolific and the negative effects of herbivory are therefore mitigated. It is for these reasons that new, potentially more damaging biocontrol agents are being considered for release. The water hyacinth grasshopper, Cornops aquaticum Brüner (Orthoptera: Acrididae), which is native to South America and Mexico, was brought into quarantine in Pretoria, South Africa in 1995. Although the grasshopper was identified as one of the most damaging insects associated with water hyacinth in its native range, it has not been considered as a biocontrol agent for water hyacinth anywhere else in the world. After extensive host-range testing which revealed it to be safe for release, a release permit for this candidate agent was issued in 2007. However, host specificity testing is no longer considered to be the only important component of pre-release screening of candidate biocontrol agents. Investigating biological and ecological aspects of the plant-herbivore system that will assist in determination of potential establishment, efficacy and the ability to build up good populations in the recipient environment are some of the important factors. This thesis is a pre-release evaluation of C. aquaticum to determine whether it is sufficiently damaging to water hyacinth to warrant its release. It investigated interactions between the grasshopper and water hyacinth under a range of nutrient conditions found in South African water bodies as well as the impact of the grasshopper on the competitive performance of water hyacinth. Both plant growth rates and the response of water hyacinth to herbivory by the grasshopper were influenced by nutrient availability to the plants. The ability of water hyacinth to compensate for loss of tissue through herbivory was greater under eutrophic nutrient conditions. However, a negative linear relationship was found between grasshopper biomass and water hyacinth performance parameters such as biomass accumulation and leaf production, even under eutrophic conditions. Water hyacinth’s compensatory ability in terms of its potential to mitigate to detrimental effects of insect feeding was dependent on the amount of damage caused by herbivory by the grasshopper. Plant biomass and the competitive ability of water hyacinth in relation to another freefloating aquatic weed species were reduced by C. aquaticum under eutrophic nutrient conditions, in a short space of time. It was also found that grasshopper feeding and characteristics related to their population dynamics such as fecundity and survival were significantly influenced by water nutrient availability and that environmental nutrient availability will influence the control potential of this species should it be released in South Africa. Cornops aquaticum shows promise as a biocontrol agent for water hyacinth but additional factors that were not investigated in this study such as compatibility with the South African climate and the current water hyacinth biocontrol agents need to be combined with these data to make a decision on its release. Possible management options for this species if it is to be introduced into South Africa are discussed.

Anredera cordifolia (Basellaceae), Madeira vine, is a perennial, semi- succulent climber native from Paraguay to southern Brazil and northern Argentina. It has a history of weediness and difficulty of control once established. In South Africa Madeira vine has a wide range and distribution with altitudes ranging from 10-1800m above sea level. Described as a transformer species, its sheer weight is capable of breaking branches off trees, causing the potential collapse of forest canopies. Chemical and mechanical control methods are expensive, labour intensive and may provide only temporary relief. A biological control programme was therefore initiated in 2003. Cf Phenrica sp. 2 (Coleoptera: Chrysomelidae: Alticinae), was field collected from A. cordifolia in Brazil, SSW of Cascavel in the Paraná Province during a survey in November 2003. Eggs are laid in groups of 16 with the average fertility rate being 89%. After going though three larval instars, the larvae pupate in the soil with the adults eclosing after a period of 17 days. The total developmental time for a generation from egg to egg ranges between 7-8 weeks. Biological traits that favour the flea beetle as a possible biological control agent include long-lived adults (up to 5 months) and multiple generations during the summer period. Both adults and larvae feed extensively on leaves and stems and although developmental rates will slow down during the winter period, no indication of a definite diapause was found under the prevailing laboratory conditions. After completing the larval no-choice trials with twenty-six plant species from 14 plant families Phenrica sp. 2 proved to be adequately host specific, as larval development was only supported by 3 Basellaceae species (including the control A. cordifolia) and one Portulacaceae species. All of these are introduced species in South Africa. The only indigenous Basella species could not be tested as it has a very marginal distribution, and because it’s inconspicuous nature, it is seldom seen or collected. Adult multi-choice trials were restricted to species that could sustain larval development to give some indication of the acceptability of these species for adult feeding and oviposition. Although adult feeding was initially concentrated on B. alba, the oviposition preference was clear-cut as females only oviposited on A. cordifolia. In order to quantify the impact of Phenrica sp. 2 on plant biomass and to assess the incidence and intensity of foliar damage, a pair of adults was confined to the host plant, for 2 generations, with different levels of larval densities. The results indicated that the host plant, due to both larval and adult feeding, suffered leaf losses of up to 55%. Anredera cordifolia was however still capable of enlarging the root mass despite suffering huge leaf losses. This would imply that A. cordifolia has an effective re-growth capacity and it will only be vulnerable to attack of the storage organs that enable re-growth, or to repeated attack of other plant parts through which reserves are exhausted. Unfortunately the period of exposure (24 days) was too short to prove that Phenrica sp. 2 impacts on the below ground dry mass, but should the plant be completely defoliated, as was observed in the field, the host plant would be forced to deplete stored resources. Phenrica sp.2 has shown to be very host specific and although A.cordifoia loses its leaves during the winter period in most provinces in South Africa, the adults are long-lived and should be able to survive the leafless periods. Further more the relatively short life cycle, high fecundity and 3 generations per year should theoretically insure a strong population build-up that would improve the chances of establishment in the field. All indications are that Phenrica sp. 2 is an agent well worth considering for the biological control of A. cordifolia.

Varieties of Lantana camara (lantana) have been introduced into many countries of the world as ornamental plants and have become invasive weeds in many countries including South Africa. In South Africa, it mostly invades the sub-tropical eastern and northern range. Mechanical and chemical control options are expensive and ineffective. A biocontrol programme was initiated in South Africa in 1961. To date, 22 insect species, and a fungus have been introduced, of these 10, and the fungus have established. Three indigenous lepidopteran species and an exotic generalist pest mealybug are also associated with the weed. The variable success of some of the agents released on L. camara worldwide has been ascribed to a few factors. One important aspect is the large range of varieties encountered in the field. It is therefore essential to be able to predict the possible establishment and impact of agents on many varieties. Laboratory trials on five of the established agents showed clear varietal preferences. In the field, most of the biocontrol agents had limited geographic ranges, linked to altitudinal conditions, as higher populations were recorded at low lying warm summer rainfall areas. A pink and orange flower corolla lobe and throat colour combination and plants with few to medium leaf hairs were most abundant in South Africa. Most of the agent species had individual preferences towards different flower colour combinations, as the agents built up different population levels on varieties in the field, within the suitable geographic region for the insect species. Eight agents preferred smooth leaved varieties, while three preferred hairy leaves, and three had no specific preference to leaf hairiness. Varietal preferences thus did play a significant role in agent populations and accompanied impact achieved in the field. New candidate agents need to be proven specific under quarantine conditions and the results extrapolated to predict specificity in the field, while avoiding potential non-target effects. Many authors have questioned the validity of laboratory host specificity trials. The conventional wisdom is that insects portray a far wider host range in the laboratory than what they would do in the field. In other words, laboratory studies measure the physiological host range of an agent and are conservative and usually don’t reflect the ecological host range of agents in the field. To avoid unnecessary rejections of biocontrol agents, this study has made a retrospective study of the host specificity of agents established in the field. Their laboratory and field host ranges were compared and it was found that virtually all the agents reflect similar or less non-target effects in the field than predicted during multiple choice trials. Of the 14 agents, only one introduced species, Teleonemia scrupulosa, and the indigenous species, Hypena laceratalis and Aristea onychote were able to sustain populations on non-target species in the field in the absence of L. camara. Insect populations on non-target species were much reduced compared to that on L. camara. Furthermore non-target effects were only recorded on plant species closely related to the target weed. The multiple choice trials therefore predict field non-target effects accurately. Predictions of non-target effects of candidate agents can therefore be accurately predicted by laboratory studies, in terms of species likely to be affected and to what extent. One field that need further study though is the impact of non-target effects, especially on Lippia species by L. camara biocontrol agents.