The inadequacy of arbitrarily classifying interactions between species as antagonistic, neutral or mutualistic has become clear in recent years. Both direct and indirect interactions between species can vary between mutualism and antagonism, depending on the intrinsic and extrinsic contexts of the interaction. This study investigated the characteristics of an ant-plant-homopteran interaction in southern Africa. The polyphagous homopteran Hilda patruelis (Tettigometridae) feeds primarily on the trunk-borne fruiting branches and figs of the Cape fig tree, Ficus sur, and produces honeydew which attracts tending ants. Ten of the sixteen ant species/species groups present on F. sur tended H. patruelis, with Pheidole megacephala the most frequent attendant. Ants attracted to F. sur by H. patruelis honeydew or other liquid food sources also preyed on insects on the tree, including adults of the small agaonid fig wasps whose larvae feed on the ovules in the developing figs. One fig wasp species (Ceratosolen capensis) is also the tree's only pollinator. No benefits to H. patruelis from being tended by ants were detected, either in terms of reduced parasitism, or predation by a lycaenid caterpillar. A P. megacephala colony foraging on a F. sur tree was found to receive a high proportion of its likely energy requirements from the tree, mainly in the form of H. patruelis honeydew, during periods when it was bearing fruit. It is probable that the H. patruelis-P. megacephala interaction constitutes a direct mutualism at times, but that benefits to the homopteran are intermittent or weak. Both H. patruelis and ants benefitted from F. sur, directly or indirectly, through the provision of food (and for some ants, nesting sites). The removal of phloem sap by H. patruelis did not detectably reduce the trees' reproductive output, either in terms of pollinator or viable seed production. The indirect effects of ant and H. patruelis presence on the F. sur trees were on average positive, as ants preyed disproportionately heavily on fig wasp species parasitic on or competing with the pollinator, thus increasing pollinator production. Effects of ant presence on seed production were not investigated, but have been demonstrated as beneficial elsewhere. However, there is great varatlon both in the composition of the wasp fauna arriving to oviposit at different crops, and in ant densities per fig, on several temporal and spatial scales. This results in high variability in the effects of ants on the pollinator and, through it, the tree, from positive to zero and potentially even negative. Despite this conditionality of beneficial outcomes for the tree, the mean effect of ants on the F. sur population studied was to increase pollinator production by up to nearly 20%. This study is among the few to have demonstrated an overall benefit to a plant of having homopteran-tending ants present on it.

Fig trees (Ficus spp., Moraceae) and fig wasps (Chalcidoidea, Agaonidae) are uniquely associated. In one fig wasp group, the pollinators (Agaoninae), each species is generally host species-specific. The relationship is one of obligate mutualism where the wasps provide pollination services and in return utilises some of the ovules for larval development. Non-pollinating fig wasps (generally belonging to subfamilies other than the Agaoninae) may be gallers or parasitoids, and can also be host species-specific. In the accompanying studies we examined the factors governing the interactions between fig wasps and their host trees. Surveys of fig trees and their associated pollinating fig wasps conducted in southern Africa, Madagascar and The Comores generally confirmed their specific relationships. An examination of F. sycomorlls in Madagascar resulted in the reclassification of F. sakalavarum as a distinct species with its own specific pollinator species. Biological and chemical evidence is presented demonstrating that the pollinators were able to distinguish their hosts through volatiles which emanated from the figs when they were ready to be pollinated. Environmental factors were found to influence wasp behaviour. Ambient temperature governed the timing of wasp emergence from their natal figs. When dispersing from their natal figs, the fig wasps flew upwards and then were blown downwind. Once nearing trees bearing figs ready to be pollinated, the wasps lost height and flew upwind towards the trees. E. baijnathi females apparently avoided figs which already contained a conspecific foundress. Scanning electron microscope studies of pollinating female fig wasp antennae showed that while all the species possessed multiporous plate sensilla, in only a few species were these sensilla elongated. Multiporous plate sensilla elongation is rare or absent among other female chalcids and may have evolved within the Agaoninae in order to facilitate their location on receptive host figs. Pollinator choice specificity appears to break down in a number of cases. In the first case examined, two pollinator species were recorded from the figs of African F. sycomorus. One. C. arabicus, pollinates the figs while the other, C. galili, acts as a 'cuckoo' by utilising some of ovules for oviposition without providing pollen. In the second case three pollinating fig wasp species were recorded from the rigs of F. lutea. Two were found to be incidental visitors and were not specifically attracted to the tree. The hybrid seeds from these crosses were successfully germinated but the seedlings did not grow passed the cotyledon stage of their development. In the concluding study the consequences of Ficus phenology and the structure of the fig's unusual inflorescence on the nonpollinating fig wasp community were examined. Various factors affecting the population levels and species richness were also examined. Future possible research directions were discussed.