The maxillary palps of Tephritidae are selectively tuned to food volatiles and diverge with ecology.

The maxillary palp is an auxiliary olfactory organ in insects, which, different from the antennae, is equipped with only a few olfactory sensory neuron (OSN) types. We postulated that these derived mouthpart structures, positioned at the base of the proboscis, may be particularly important in mediating feeding behaviors. As feeding is spatio-temporally segregated from oviposition in most Tephritidae, this taxonomic group appears quite suitable to parse out sensory breadth and potential functional divergence of palps and antennae. Scanning electron microscopy and anterograde staining underlined the limited palpal olfactory circuit in Tephritidae: only three morphological subtypes of basiconic sensilla were found, each with two neurons, and project to a total of six antennal lobe glomeruli in Bactrocera dorsalis. Accordingly, the palps detected only few volatiles from the headspace of food (fermentation and protein lures) and fruit (guava and mango) compared to the antennae (17 over 77, using gas-chromatography coupled electrophysiology). Interestingly, functionally the antennae were more tuned to fruit volatiles, detecting eight times more fruit than food volatiles (63 over 8), whereas the number of fruit and food volatile detection was more comparable in the palps (14 over 8). As tephritids diverge in oviposition preferences, but converge on food substrates, we postulated that the receptive ranges of palpal circuits would be more conserved compared to the antennae. However, palpal responses of three tephritid species that differed in phylogenetic relatedness and ecologically niche, diverged across ecological rather than phylogenetic rifts. Two species with strongly overlapping ecology, B. dorsalis and Ceratitis capitata, showed inseparable response profiles, whereas the cucurbit specialist Zeugodacus cucurbitae strongly diverged. As Z. cucurbitae is phylogenetically placed between B. dorsalis and C. capitata, the results indicate that ecology overrides phylogeny in the evolution of palpal tuning, in spite of being predisposed to detecting food volatiles.

Dispersal and competitive release affect the management of native and invasive tephritid fruit flies in large and smallholder farms in Ethiopia.

African horticulture is seriously affected by fruit flies, both native and invasive. Novel sustainable control methods need testing against the backdrop of smallholder-dominated farming of Africa. We evaluated the potential of male-specific attractants (parapheromones) laced with insecticide to suppress the alien invasive Bactrocera dorsalis and native Ceratitis capitata. In large-scale guava, methyl-eugenol (ME)-bait stations combined with toxic protein baits suppressed B. dorsalis within 8 months but resulted in a resurgence of the displaced Ceratitis capitata. In smallholder farms, intervention using SPLAT-ME laced with spinosad was surprisingly unsuccessful. Subsequent mark-release-recapture experiments showed high dispersal rates of flies, covering many times a typical farm size, leading to a continuous influx of flies from surrounding areas. Several other factors important for intervention were evaluated. SPLAT-MAT-ME dollops remained attractive for over two weeks, although gradually becoming less attractive than fresh baits. Further, competitive displacement was observed: C. capitata selectively emerged from fruits in which B. dorsalis infestation was low. Finally, we evaluated whether ME could be combined with C. capitata male attractants [trimedlure (TML) and terpinyl acetate (TA)] without affecting attraction. Combining male lures did not affect catches directly, although at very high populations of B. dorsalis attracted to ME interfered with C. capitata trap entry. Although ME-based methods can effectively suppress B. dorsalis, they were not effective at single smallholder scale due to the high dispersive propensity of tephritids. Further, competitive release implies the need for a combination of lures and methods. These observations are important for developing control schemes tailored for African smallholder settings.

Detection of Volatile Constituents from Food Lures by Tephritid Fruit Flies.

Tephritid fruit flies require protein for sexual and gonotrophic development. Food-based lures are therefore widely used in strategies to detect and control fruit flies in the Tephritidae family. However, these baits are attractive to a broad range of insect species. We therefore sought to identify volatiles detected by the fly antennae, with the goal to compose lures that more specifically target tephritids. Using gas chromatography-coupled electroantennographic detection (GC-EAD) we screened for antennal responses of four important tephritid species to volatile compounds from five commercially available protein-based baits. Antennal active compounds were reconstituted in synthetic blends for each species and used in behavioral assays. These species-based blends were attractive in olfactometer experiments, as was a blend composed of all antennally active compounds from all the four species we observed (tested only in Bactrocera dorsalis, Hendel). Pilot field tests indicate that the blends need to be further evaluated and optimized under field conditions.