An assessment of the anatomical costs of extremely long proboscid mouthparts can contribute to the understanding of the evolution of form and function in the context of insect feeding behaviour. An integrative analysis of expenses relating to an exceptionally long proboscis in butterflies includes all organs involved in fluid feeding, such as the proboscis plus its musculature, sensilla, and food canal, as well as organs for proboscis movements and the suction pump for fluid uptake. In the present study, we report a morphometric comparison of derived long-tongued (proboscis approximately twice as long as the body) and short-tongued Riodinidae (proboscis half as long as the body), which reveals the non-linear scaling relationships of an extremely long proboscis. We found no elongation of the tip region, low numbers of proboscis sensilla, short sensilla styloconica, and no increase of galeal musculature in relation to galeal volume, but a larger food canal, as well as larger head musculature in relation to the head capsule. The results indicate the relatively low extra expense on the proboscis musculature and sensilla equipment but significant anatomical costs, such as reinforced haemolymph and suction pump musculature, as well as thick cuticular proboscis walls, which are functionally related to feeding performance in species possessing an extremely long proboscis.

Few species of true butterflies (Lepidoptera: Papilionoidea) have evolved a proboscis that greatly exceeds the length of the body. This study is the first to examine the morphology of an extremely long butterfly proboscis and to describe how it is used to obtain nectar from flowers with very deep corolla tubes. The proboscis of Eurybia lycisca (Riodinidae) is approximately twice as long as the body. It has a maximal length of 45.6 mm (mean length 36.5 mm ± 4.1 S.D., N = 20) and is extremely thin, measuring only about 0.26 mm at its maximum diameter. The proboscis has a unique arrangement of short sensilla at the tip, and its musculature arrangement is derived. The flower handling times on the preferred nectar plant, Calathea crotalifera (Marantaceae), were exceptionally long (mean 54.5 sec ± 28.5 S.D., N = 26). When feeding on the deep flowers remarkably few proboscis movements occur. The relationship between Eurybia lycisca and its preferred nectar plant and larval host plant, Calathea crotalifera, is not mutualistic since the butterfly exploits the flowers without contributing to their pollination. We hypothesize that the extraordinarily long proboscis of Eurybia lycisca is an adaptation for capitalizing on the pre-existing mutualistic interaction of the host plant with its pollinating long-tongued nectar feeding insects.

Extremely long proboscides are rare among butterflies outside of the Hesperiidae, yet representatives of several genera of skipper butterflies possess proboscides longer than 50 mm. Although extremely elongated mouthparts can be regarded as advantageous adaptations to gain access to nectar in deep-tubed flowers, the scarcity of long-proboscid butterflies is a phenomenon that has not been adequately accounted for. So far, the scarceness was explained by functional costs arising from increased flower handling times caused by decelerated nectar intake rates. However, insects can compensate for the negative influence of a long proboscis through changes in the morphological configuration of the feeding apparatus. Here, we measured nectar intake rates in 34 species representing 21 Hesperiidae genera from a Costa Rican lowland rainforest area to explore the impact of proboscis length, cross-sectional area of the food canal and body size on intake rate. Long-proboscid skippers did not suffer from reduced intake rates due to their large body size and enlarged food canals. In addition, video analyses of the flower-visiting behaviour revealed that suction times increased with proboscis length, suggesting that long-proboscid skippers drink a larger amount of nectar from deep-tubed flowers. Despite these advantages, we showed that functional costs of exaggerated mouthparts exist in terms of longer manipulation times per flower. Finally, we discuss the significance of scaling relationships on the foraging efficiency of butterflies and why some skipper taxa, in particular, have evolved extremely long proboscides.

Extremely long proboscides of insect flower visitors have been regarded as an example of a coevolutionary arms race, assuming that these insects act as efficient pollinators for their nectar host plants. However, the effect of proboscis length on generalized or specialized flower use remains unclear and the efficiency of butterfly pollination is ambiguous. Neotropical Hesperiidae feature a surprising variation of proboscis length, which makes them a suitable study system to elucidate the role of extremely long-proboscid insects in plant-pollinator networks. The results of this study show that skippers with longer proboscides visit plant species with deep-tubed flowers to take up food, but do not pollinate them. Skippers equipped with extremely long proboscides seldom include short-tubed flowers in their diet nor visit more plant species than those with shorter proboscides. Our observations indicate that the extremely long-proboscid skippers steal nectar from their preferred nectar host plants, Calathea sp., instead of contributing to their pollination. Finally, we discuss the impact of nectar robbery by these butterflies on their nectar host plants and their legitimate pollinators, euglossine bees.

Social wasps perform a variety of tasks with their mouthparts. Female workers use them to feed on carbohydrate-rich fluids, to build nests by collecting wood fibers and forming paper, to hunt and manipulate insect prey for feeding larvae as well as for brood care. Since male wasps neither feed on insects nor participate in nest building, sex-specific differences in mouthpart morphology are expected. Despite these different applications, general mouthpart morphology of male and female wasps from the genus Vespula was similar. However, males possessed significantly shorter mandibles with fewer teeth than females. Furthermore, the adductor muscles of the mandibles were distinctly smaller in males than in females. Male wasps showed a higher number of sensilla on the mandibles and the labial palpi. Mouthpart dimorphism and functional morphology of fluid uptake are discussed.