Revisiting trends in morphology of antennal sensilla in scarabaeoid beetles

Phytophagous scarab beetles associated with angiosperms have characteristically enlarged lamellate antennae and exhibit a striking morphological variation of sensilla. In this study, we compared the morphology of antennal surface of 62 species Scarabaeoidea using SEM microscopy, particularly also in light of their evolution in association with angiosperms. We investigated the correlation of antennal sensilla morphology, i.e., their structure and distribution, with species diversity and lineage diversification rates. A high diversity of sensilla was observed but also multiple transitional forms, even on the same antennomere. We interpreted this as evidence for a high evolutionary plasticity. We recognized clear patterns of convergence and repeated evolution of certain types of placoid sensilla. One main tendency found in the phytophagous Pleurostict chafers was a shift from sensilla trichodea to placoid-like sensilla, apparently also enhanced by the increase of the lamellate antennal surface, either by size or number of the lamellae. This trend occurred not only in the Pleurosticts, but also in Glaphyridae, a second angiosperm-associated lineage of Scarabaeoidea. However, our results suggest no direct relation between species diversity or the rate of diversification and general sensilla morphology, i.e., the origin of placoid sensilla. This could be explained not only by species-poor lineages also possessing placoid sensilla but also by otherwise successful and species rich groups having sensilla trichodea (e.g., dung beetles). Results further reveal the need to refine current phylogenetic hypotheses by more comprehensive taxon sampling and to expand the molecular characterization of pheromones and odor binding proteins to better understand the role of chemical communication in scarab diversification.