Lantana camara Linnaeus (Verbenaceae), commonly known as lantana, is a highly invasive weed in many parts of the world. In South Africa it is naturalized in several provinces where it invades pastures, riverbanks, mountain slopes and valleys and commercial and natural forests, forming dense, impenetrable thickets. Chemical and mechanical control methods are expensive, labour intensive and provide only temporary relief as cleared areas are rapidly reinfested by seedlings and coppice growth. A biological control programme was initiated in South Africa in the 1960s, but despite the establishment of 11 agent species, it was considered to have had limited success. Several factors are thought to restrict the impact of the biocontrol agents. Firstly, L. camara occurs in a range of climatic regions, some of which are unsuitable for the establishment of agent species of tropical and subtropical origin. Secondly, L. camara is the result of hybridization between several Lantana species, forming a complex of hybridized and hybridizing varieties in the field, which match none of the Lantana species in the region of origin. This causes partial insect-host incompatibility, displayed as varietal preference. Thirdly, parasitism appears to have significantly reduced the effectiveness of several natural enemies. In spite of all these constraints, biological control has reduced invasion by L. camara by 26%. However, the weed is still very damaging and additional natural enemies are required to reduce infestations further. A flea-beetle species, Alagoasa extrema Jacoby (Coleoptera: Chrysomelidae), was collected from several sites in the humid subtropical and tropical regions of Mexico, and imported into quarantine in South Africa and studied as a potential biocontrol agent for L. camara. Favourable biological characteristics of this beetle included long-lived adults, several overlapping generations per year, and high adult and larval feeding rates. Observations from the insect’s native range and studies in South Africa suggest that A. extrema would probably be more suited to the subtropical, rather than the temperate areas in South Africa. Laboratory impact studies indicated that feeding damage by A. extrema larvae, over a period spanning the larval stage (16 to 20 days), reduced the above-ground biomass of L. camara plants by up to 29%. Higher larval populations resulted in a higher reduction of biomass. Varietal preference and suitability studies indicated that A. extrema exhibits a degree of varietal preference under laboratory conditions, with one of the white pink L. camara varieties proving the most suitable host. This variety is one of the most damaging varieties in South Africa and is particularly widespread in Mpumalanga Province. Although A. extrema proved to be damaging to L. camara, laboratory host range trials showed it to be an oligophagous species, capable of feeding and developing on several non-target species, especially two native Lippia species (Verbenaceae). The host suitability of these species was marginally lower than that of L. camara and the potential risk to these indigenous species was deemed to be too high to warrant release. It was therefore recommended that A. extrema not be considered for release in South Africa.

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.

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.

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.