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.

Interactions between insects have been shown to be important regulators of population abundances and dynamics as well as drivers of spatial segregation and distribution. These are important aspects of the ecology of insects used in biological control and may have implications for the overall success of a particular programme. In the history of biological control there has been a tendency to release a suite of agents against a weed, which in some cases has increased the level of success, while in others little change has been observed. In most of these cases the implications of increasing the level of complexity of the system is not taken into account and there is little research on the effect of releasing another agent into the system. A brief meta-analysis was done on all the biological control programmes initiated in South Africa. Emphasis was placed on multi-species releases and the effects that overlapping niches were having on the number of agents responsible for the success of a programme. Where overlapping niches were present among agents released the number of agents responsible for success was lower than the number established. Water hyacinth, Eichhornia crassipes (Martius) Solms-Laubach in South Africa has more arthropod agents released against it than anywhere else in the world, yet control has been variable. If the biology and host utilisation of all the agents against water hyacinth is considered, a definite overlap of niches is apparent in at least one life stage of all the agents. Therefore the probability of these insects interacting is high, especially if they are established at the same site in the field. Three of the insects released in South Africa have been selected to investigate possible interactions. They are Neochetina eichhorniae Warner, Neochetina bruchi Hustache and Eccritotarsus catarinensis (Carvalho). Y-tube olfactometer bioassays were used to measure responses of these insects to water hyacinth with prior feeding damage by either conspecifics or heterospecifics. This was done to determine whether olfactory cues played a role in host acceptability and avoidance of conspecifics or heterospecifics. The insects were given a choice between damaged and undamaged plants in various combinations. There was a significant preference for the undamaged plants when given a choice between undamaged and damaged plants. However when the insects were given a choice between two damaged plants there was no discrimination between heterospecific or conspecific damaged plants. This may indicate that there is little or no ecological cost for the insect to share a plant with other insects utilising a similar resource. Insect – insect interactions were investigated in a common garden plot experiment to measure the impact that pairwise combinations of the insect may have on their performance. There was a significant interaction between the mirid E. catarinensis and the weevil N. eichhorniae, with the weevil not performing as well when in combination with the mirid than when alone. Interestingly there was a negative interaction between the two weevil species when in combination, however it was impossible to determine which species was being affected if not both. None of the insects performed significantly better when in combination with another insect. A field study on Wriggleswade Dam in the Eastern Cape, South Africa was initiated to determine whether the relationship between the mirid E. catarinensis and the weevil N. eichhorniae could be determined in the field. The performance of the insects at the different sites in the field suggests that there was an interaction between the agents. This interaction did not limit the establishment of either insect at a site, but it did result in one insect dominating at a site over another. Interactions between the three species of insect tested in this thesis suggest that there are both negative and neutral relationships between them. A basic comparison between the insect performances from 15 sites around the country was done to determine if the spatial segregation observed in the field could be extrapolated to the natural South African situation. The interaction observed between N. eichhorniae and E. catarinensis does seem to extrapolate to the general South African situation where there is definite spatial segregation on a landscape level. The co–occurrence of the two Neochetina weevils at these sites suggests that the negative relationship observed between them in the common garden experiment does not extrapolate to the field. The results from this thesis suggest that the interactions between the agents tested would not limit establishment or have significant ramifications on performance. However, there may be spatial and temporal segregation of these species in the introduced range.

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.