The Autecology of Bactrocera cacuminata (Hering) (Diptera:Tephritidae:Dacinae): Functional Significance of Resources

Abstract

This thesis investigated the autecology of the dacine species, Bactrocera cacuminata (Hering) (Diptera: Tephritidae: Dacinae). I specifically focused on the adult phase of the life cycle and resources believed to be significant to this life stage. The prevailing paradigm in dacine ecology predicts that the larval host plant serves as the centre of dacine activity, a state mediated by mutualistic associations with fruit fly-type bacteria. Contrary to predictions, an explicit test of this hypothesis found that the host plant of B. cacuminata, Solanum mauritianum Scopoli, acted almost exclusively as a site for oviposition and larval development. Other key adult behaviours, most notably feeding and mating, were rare at the host plant. Even in disturbed habitats, the paucity of key adult behaviours such as mating was striking. Adult flies of this species were therefore hypothesized to be utilizing other components of their habitat, i.e. resources vital to their life history requirements. Some of the resources that B. cacuminata are known to respond to include sugar, protein, methyl eugenol and the host plant. The latter three resources are believed to be critical in the reproductive success of dacine flies in general. I assessed the physiological status of flies arriving at these resources to determine if flies of different status foraged for resources differently. In dacines, the internal reproductive structures of the male and female flies have been used as predictors of physiological status. I quantified expansion of the male ejaculatory apodeme in B. cacuminata with age of fly and found that there is a threshold apodeme size that is strongly correlated Abstract with sexual maturity. Maturity of female flies could be accurately predicted by ovarian development. Using these methods to assess the physiological and nutritional status of flies arriving at resources (larval host plant, protein and methyl eugenol) in the field, I discovered that only sexually mature and mated females were responding to the host plant, while the males at the host plant were sexually immature. This confirmed the hypothesis that the host plant primarily served as an oviposition site. Additionally, this study revealed that sexually mature males with high nutritional reserves were most commonly collected at methyl eugenol (a plant-derived chemical that elicits a strong response in males of many dacine species) at dusk, the time of peak sexual activity in this species. This indicated that methyl eugenol was perhaps a significant resource in the context of the reproductive behaviour of this species. Methyl eugenol (ME) is one of group of phenyl propanoids to which males of certain species of Dacinae respond. The current hypothesis of the role of these phenyl propanoids is that they function as pheromone precursor chemicals. Response to these chemicals is hypothesized to be a trait under sexual selection. In Bactrocera dorsalis (Hendel), this effect is so strong that a single feeding on ME results in a strong mating advantage up to a month after males feed on the chemical. Bactrocera cacuminata fed on multiple occasions on ME in a laboratory bioassay. After a single 24-hour exposure to ME, investigations of mating competitiveness did not reveal any obvious advantage for ME-fed males over unfed males. However, ME-fed males did enjoy a higher mating success 16 and 32 days after exposure to the chemical, suggesting that some physiological benefits unrelated to the pheromone synthesis was driving this delayed advantage. Investigation of the physiological consequences of feeding on ME revealed no enhancement of nutritional or energetic reserves, suggesting that the delayed mating advantage observed was more likely a chance event. An alternate hypothesis about the proximate function of ME, proposed by Robert Metcalf, is that it serves as a mate rendezvous site. As mating behaviour was notably absent at the host plant, I tested Metcalf’s hypothesis. A field-cage experiment, spatially separating adult resources (host plant, methyl eugenol, sugar and protein) clearly demonstrated that methyl eugenol was functioning as a mate rendezvous stimulus for B. cacuminata. This is the first direct support for Metcalf’s hypothesis. A synthesis of the literature revealed that significantly greater ecological and evolutionary information was required to understand the basis of dacine response to phenyl propanoids. Different dacine species may be utilizing these chemicals differently, even if their evolutionary origin may have been as a plant based kairomone. My studies show that generalizations on the ecology and behaviour of Dacinae, often extrapolated from research on a few pest species, do not hold up in the case of B. cacuminata. This suggests that a more autecological, species-specific approach is required in dacine research, before any predictive generalizations can be made.