Mixed xylem and phloem sap ingestion in sheath-feeders as normal dietary behavior: Evidence from the leafhopper...

In phytophagous piercing-sucking insects, salivary sheath-feeding species are often described as xylem- or phloem-sap feeding specialists. Because these two food sources have very different characteristics, two feeding tactics are often associated with this supposed specialization. Studying the feeding behavior of insects provides substantial information on their biology, ecology, and evolution. Furthermore, study of feeding behavior is of primary importance to elucidate the transmission ability of insects that act as vectors of plant pathogens. In this study, we compared the durations of ingestion performed in xylem versus phloem by a leafhopper species, Scaphoideus titanus Ball, 1932. This was done by characterizing and statistically analyzing electrical signals recorded using the electropenetrography technique, derived from the feeding behaviors of males and females. We identified three groups of S. titanus based on their feeding behavior: 1) a group that reached the phloem quickly and probed for a longer time in phloem tissue than the other groups, 2) a group that reached the xylem quickly and probed for a longer time in xylem tissue than the other groups, and 3) a group where individuals did not ingest much sap. In addition, the numbers and durations of waveforms representing ingestion of xylem and phloem saps differed significantly depending on the sex of the leafhopper, indicating that the two sexes exhibit different feeding behaviors. Males had longer phloem ingestion events than did females, which indicates that males are greater phloem feeders than females. These differences are discussed, specifically in relation to hypotheses about evolution of sap feeding and phytoplasma transmission from plant to plant.