Evolutionary functional morphology of the proboscis and feeding apparatus of hawk moths (Sphingidae: Lepidoptera).

The morphology of the proboscis and associated feeding organs was studied in several nectar-feeding hawk moths, as well as a specialized honey-feeder and two supposedly nonfeeding species. The proboscis lengths ranged from a few millimeters to more than 200 mm. Despite the variation in proboscis length and feeding strategy, the principle external and internal composition of the galeae, the stipes pump, and the suction pump were similar across all species. The morphology of the smooth and slender proboscis is highly conserved among all lineages of nectar-feeding Sphingidae. Remarkably, they share a typical arrangement of the sensilla at the tip. The number and length of sensilla styloconica are independent from proboscis length. A unique proboscis morphology was found in the honey-feeding species Acherontia atropos. Here, the distinctly pointed apex displays a large subterminal opening of the food canal, and thus characterizes a novel type of piercing proboscis in Lepidoptera. In the probably nonfeeding species, the rudimentary galeae are not interlocked and the apex lacks sensilla styloconica; galeal muscles, however, are present. All studied species demonstrate an identical anatomy of the stipes, and suction pump, regardless of proboscis length and diet. Even supposedly nonfeeding Sphingidae possess all organs of the feeding apparatus, suggesting that their proboscis rudiments might still be functional. The morphometric analyses indicate significant positive correlations between galea lumen volume and stipes muscle volume as well as the volume of the food canal and the muscular volume of the suction pump. Size correlations of these functionally connected organs reflect morphological fine-tuning in the evolution of proboscis length and function.

Morphological fine tuning of the feeding apparatus to proboscis length in Hesperiidae (Lepidoptera).

The form and function of the hesperiid feeding apparatus was studied in detail. The butterflies in the family Hesperiidae are of particular interest because the longest proboscis ever recorded in Papilionoidea was found in the Neotropical genus Damas. We focused on the functional morphology by comparing proboscis morphology as well as size and composition of both the stipes pump and the cibarial suction pump in skippers with short and extremely long proboscis. Results revealed that all studied Hesperiidae have the same proboscis micromorphology and sensilla endowment regardless of the proboscis length. However, the numbers of internal muscles of the proboscis, the morphology of the stipes pump as well as the pumping organs for nectar uptake are related to the proboscis length. We conclude that the low number of tip sensilla compared to proboscis length is responsible for remarkably longer manipulation times of long-proboscid species during flower visits. The organs for proboscis movements and nectar uptake organs are well tuned to the respective proboscis length and are accordingly bigger in species with a proboscis that measures twice the body length.

Mouthparts and nectar feeding of the flower visiting cricket Glomeremus orchidophilus (Gryllacrididae).

Glomeremus orchidophilus (Gryllacrididae) is a flower visiting cricket on the tropical island La Réunion. This species is the only Orthoptera shown to be a pollinator of a plant. We studied its nectar feeding behavior and mouthpart morphology in detail. Since G. orchidophilus possesses biting-and-chewing mouthparts, our objective was to find behavioral and/or structural specializations for nectar-feeding. The comparative analysis of feeding behavior revealed that fluid is taken up without movements of the mouthparts in Glomeremus. A comparative morphological examination of two Glomeremus species, together with several representatives of other Gryllacrididae and other Ensifera taxa revealed subtle adaptations to fluid feeding in Glomeremus. All representatives of Gryllacrididae were found to possess a distinct patch of microtrichia at the tip of their galeae. However, in Glomeremus a channel is formed between the distal components of the maxillae and the mandibles on each side of the body. Micro-CT and SEM examination revealed a longitudinal groove that extends over the galea beginning at the patch of microtrichia in the studied Glomeremus species. We hypothesize that the microtrichia take up fluid by capillarity and the action of the cibarium and pharyngeal pumps transports fluid along the channels between the maxillae and mandibles into the preoral cavity. These mouthpart features allow nectar uptake from flowers that is unique in Orthoptera.

The extremely long-tongued neotropical butterfly Eurybia lycisca (Riodinidae): proboscis morphology and flower handling.

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.