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.

The biological control of the weedy complex Lantana camara (L.) (Verbenaceae) has been ongoing in South Africa for over 40 years. Despite this, the weed is still not under sufficient control and continues to invade new territories in the country. The biological control programme needs to be bolstered with releases of new and potentially damaging biological control agents. A promising biological control agent endemic to Central America, Falconia intermedia (Distant) (Hemiptera: Miridae), was imported into quarantine from Jamaica in 1994. This agent was released on sites in KwaZulu-Natal and Limpopo provinces of South Africa in 1999. Even though it initially established and damaged L. camara, populations died out at most of the release sites. As varietal difference and adverse climate have been cited as the reason for non-establishment and ineffective control in L. camara biocontrol programmes worldwide, this study attempts to investigate the role that these two factors play in this weed herbivore relationship. Laboratory no-choice trials were conducted to determine the varietal performance of F intermedia, among five Eastern Cape varieties of the weed from East London, Whitney Farm, Heather Glen Farm, Port Alfred and Lyndhurst Farm, and a variety from the Plant Protection Research Institute (PPRI), Pretoria. However, there were differences in performance as the adult mirids performed better on white-pink varieties from Whitney Farm and Heather Glen Farm. To test varietal preference in field conditions, field releases of F intermedia were also made at East London, Whitney Farm, Heather Glen Farm, Port Alfred and Lyndhurst Farm. Post release evaluations were conducted monthly for two years (2002 and 2003). The insect established at East London and Whitney Farm, both of which have white-pink varieties. Insect populations quickly died out at the Lyndhurst Farm and Port Alfred sites, which have dark pink varieties. It is suggested that field conditions may have resulted in poor plant quality and led indirectly to varietal preference, and to non-establishment at these two sites. With the onset of cooler weather, populations disappeared at Heather Glen Farm. This suggested that F. intermedia was suitable for release in more subtropical areas within South Africa where climatic conditions would be suitable throughout the year. The mirid performed well at Whitney Farm, resulting in significant reduction in plant growth parameters such as height and percentage cover, and increasing the cover of other flora growing beneath L. camara plants. Finally, ways to improve the efficacy of this agent are considered in an effort towards better control of L. camara in South Africa.

Ph.D. (Zoology) / Scale insects, plant pests of quarantine importance, with specialised anatomy and unresolved phylogenetic relationships, are responsible for major economic losses to South Africa and its trading partners. These losses may reach critical levels if the pests are not timely identified and controlled. They are currently identified based on published keys of adult females, a process that takes three days to two weeks depending on the family and the life stage of interception. In addition, agricultural commodities are often contaminated with different life stages, males or damaged specimen of these pests, making identification difficult or impossible. As a result, shipments of agricultural produce are often rejected and trade disrupted. Furthermore, pest invasions do not only occur by importation via formal channels. At times pests cross boarders as contaminants of undeclared material and may again spread on their own as they naturally expand their range. This expansion may be negatively or positively influenced by other factors such as climate change. Resolving the challenges associated with identification, phylogenetic relationships and the limited knowledge of the effects of climate change on distribution range of scale insects are the main goals of this study. Specifically (i) the development of a rapid method of species identification, (ii) the relationship between and within three major scale insect families the Coccoidea, Diaspididae and Pseudococcidae and (iii) the effect of climate change on the future distribution range of scale insects in South Africa were explored...

Lantana camara L. (Verbenaceae) remains a highly invasive and ecologically damaging weed in South Africa, despite some 50 years of biological control efforts. Lack of success has been ascribed to varietal differences, climate and predation of agents but these have not been tested. In this study, the effects of ant predation were tested on populations of three biological control agents for L. camara. Colonies of two species, Crematogaster sp. 1 and 2 were investigated. Crematogaster sp. 1 colonies were offered no choice between immature stages of the agents Hypena laceratalis Walker (Lepidoptera: Noctuidae), Falconia intermedia Distant (Hemiptera: Miridae) or Teleonemia scrupulosa Stål (Hemiptera: Tingidae) on lantana shoots. Density-dependent predation on F. intermedia and T. scrupulosa nymphs on lantana shoots was tested using Crematogaster sp. 2 colonies. In choice experiments Crematogaster sp. 2 colonies were offered F. intermedia or T. scrupulosa nymphs on potted lantana plants. Preliminary food trials confirmed that colonies foraged for protein, thereby validating results of no-choice experiments. Crematogaster sp.1 foragers removed 50% of F. intermedia nymphs, followed by 45% of H. laceratalis larvae and only 9% of T. scrupulosa nymphs. Foragers recruited most actively to H. laceratalis larvae and significantly more H. laceratalis biomass was removed than either F. intermedia or T. scrupulosa. A trade-off existed in prey size selection because larger larvae provided considerably more biomass but required forager cooperation and a longer time to subdue than did smaller prey. This increases both forager energy expense and mortality risk by other predators. This study showed that all Crematogaster sp. 1 colonies removed small (≤10mm) H. laceratalis larvae more frequently than larvae larger than 10mm. Thus, of these biological control agents, predators probably prefer small H. laceratalis larvae. Significantly more F. intermedia than T. scrupulosa nymphs were removed by Crematogaster sp. 1, while Crematogaster sp. 2 colonies removed comparable numbers of both agent species. Falconia intermedia nymphs' fast movement triggered a predatory response by these ant species. In contrast, the relatively immobile behaviour of T. scrupulosa nymphs was identified as a highly effective predator avoidance strategy. Since T. scrupulosa nymphs are unable to escape predators by moving, they appear to depend on the presence of alternative prey attracting predator attention. At high agent and/or forager density, T. scrupulosa nymphs attempted escape, but foragers identified them as prey once they moved and caught them. Predation on F. intermedia was also density dependent in that at high nymph and/or forager densities, escape routes were congested and nymphs were more easily caught. Survival of F. intermedia and T. scrupulosa nymphs in particular was low on ant-accessed shrubs in choice experiments and high on ant-excluded shrubs. It is likely that ants significantly depress F. intermedia populations in the field since besides predation, ant foragers probably interrupt F. intermedia feeding and ovipositioning. The combination of parasitism and predation on early instar larvae may explain why H. laceratalis occurs across lantana's range in South Africa but populations remain low. It is unlikely that T. scrupulosa nymphs are habitually preyed on by ant species unless they attract attention by being mobile. Although biological control of L. camara is influenced by climate and physiological defence mechanisms, this study has shown that predation by two ant species severely impacts leaf-feeding agents for L. camara. Thus, it is recommended that future selection of additional agents to control lantana should exclude leaf-feeding .

A highly variable invasive shrub, Lantana camara L. (Verbenaceae), has been notoriously difficult to control thus far despite a well established biological control programme in South Africa. A promising leaf-feeding biological control agent, Falconia intermedia (Distant) (Hemiptera: Miridae), released to control this invasive plant eventually crashed at three out of five sites in the Eastern Cape Province. In the Mpumalanga Province, after initially colonising and building up high numbers on the L. camara stands the agent populations crashed. Several reasons for these population crashes have been suggested, but induced plant defences have not been investigated. Although plants face the challenge of herbivory by various organisms while remaining immobile, some plants may possess the ability to induce physical and/or chemical defensive responses following feeding and thus prevent further plant tissue damage and loss. Laboratory trials were conducted to determine the existence, nature and effect of physical and chemical feeding-induced responses of L. camara on the performance of the leaf-feeding biological control agent, F. intermedia. Lantana camara plants used in the study were obtained from five localities in the Eastern Cape Province, South Africa, while the insect culture was established from field populations. Plants from all varieties on which F. intermedia was released significantly increased the toughness of their leaves compared to control treatment plants. In addition, plants from three localities: Lyndhurst Farm, East London and Port Alfred, significantly increased trichome density after prolonged feeding by F. intermedia. On the three varieties showing increases in these two factors (i.e. leaf toughness and trichome density), oviposition, survival and feeding damage by the mirid agent was significantly lower on previously damaged plants. A significant negative correlation between trichome density and population numbers was found (R²= 0.52, p < 0.0003), suggesting that an increase in trichome density strongly contributes to a reduction in F. intermedia's growth. The growth and reproduction of the resistant plants was not significantly impacted by F. intermedia feeding. The defensive responses were found to be plant systemic and rapidly induced as they were elicited and expressed throughout the plant in both damaged and undamaged leaves within five weeks after insect release. Leaf toughness and trichome density were not significantly increased after feeding on plants from Whitney Farm and Heather Glen Farm. On the contrary, mirid individuals performed significantly better on plants from Whitney Farm and Heather Glen Farm than on plants of other varieties, indicating their susceptibility and suitability to the agent and the lack of induced resistance against the agent. Plants from all localities besides East London showed some level of tolerance and overcompensated for feeding damage by increasing plant growth and reproductive factors on plants fed upon. This was however only significant in two variables of the more susceptible localities, Whitney Farm and Heather Glen Farm. This increase in plant fitness did however indicate an induced defence response by these plants to feeding, a response designed to lessen the effects of agent feeding. Headspace volatile analysis was used to investigate any volatile chemical responses by L. camara due to F. intermedia feeding at two of the five localities chosen: East London and Whitney Farm. There was no significant difference in headspace volatiles emitted by leaves of plants from the East London insect infested and control treatment plants. On the Whitney Farm damaged plants however there was a 2.5 fold increase in the emission intensity of one of the three main compounds, later identified as Beta-caryophyllene. Three major chemical constituents which were found to be common to leaf volatiles of the two varieties were identified through gas chromatography-mass spectrometry (GC-MS) from the damaged and undamaged leaves of these two varieties. The methods used in collecting leaf volatiles were shown to be significant in the strength of chromatogram peaks. Using general authentication methods and purified standards, one of these was identified as the sesquiterpene, Beta-caryophyllene (C₁₅H₂₄). This compound is one of the major constituents found in isolations of L. camara varieties worldwide. This is the first such work done on a variety of L. camara in South Africa, and hopefully the beginning of more in-depth studies of the volatile organic chemicals from the numerous naturalised varieties of L. camara. It is suggested that the sum of these responses may play a role bigger than is currently understood in this plant-insect relationship. It is also argued that feeding induced plant defences may play an important role in attempts to control alien plants using insect agents.