Proboscis morphology and food preferences in nymphalid butterflies (Lepidoptera: Nymphalidae)

Many species of nymphalid butterflies only exceptionally visit flowers and feed instead on tree sap, juice of rotting fruits and other decaying substances. To investigate whether the proboscis morphology of these non-flower-visiting Nymphalidae differs from that of nectarivorous butterflies, representatives from 64 nymphalid species with known feeding preferences were examined. Morphometric comparison of the proboscis revealed characteristic differences in proboscis length, tip-region length, wall composition, and number and shape of proboscis sensilla between these two feeding guilds. The investigated non-flower-visiting species belonging to Apaturinae, Limenitidinae, Morphinae, Brassolinae, Nymphalinae and Satyrinae, possess a relatively short and light-coloured proboscis which has a long tip-region with a great number of club-shaped sensilla styloconica. Densely arranged, these sensilla form a flat brush located laterally from the openings into the food canal on the dorsal side of the tip-region. Among the non-flower-visiting species, a second type of proboscis was found in fruit-feeding Charaxinae the stout tip-region of which is equipped with more widely spaced sensilla styloconica. The investigated flower-visiting Heliconiinae, Nymphalinae, Satyrinae, Danainae and Ithomiinae are characterized by a slender, darker-coloured proboscis with a rather short tip-region bearing fewer sensilla styloconica in a loose arrangement. Discriminant analysis revealed that the tip-region length, the number of sensilla styloconica and the relative proboscis length are the most important discriminating variables between the flower-visiting and the non-flower-visiting species. The proboscis morphology of nymphalid butterflies corresponds with certain feeding habits and allow us to make predictions on their food preferences. The 'brush-tipped' proboscis seems to have a functional role in the accumulation of fluid and the uptake of liquid from wet surfaces such as rotting fruits or tree sap. We conclude from the phylogeny of the examined taxa that this derived proboscis lip morphology evolved several times independently as an adaptation to the exploitation of new food resources.