Why does the larval integument of some sawfly species disrupt so easily?: The harmful hemolymph hypothesis

The larvae of several sawfly species belonging to the Tenthredinidae (Hymenoptera) have such a low mechanical resistance in the integument that slight mechanical damage to the integument is enough to provoke the release of hemolymph at a given spot. We quantified this phenomenon, which we call "easy bleeding", by measuring the pressure needed to pierce dissected sawfly integument. We also investigated the feeding deterrence of ethanolic extracts of the hemolymph by laboratory bioassays using Myrmica rubra ant workers. These traits, integument resistance and hemolymph deterrence, were inversely related, considering 22 ten-thredinid species. A negative correlation was obtained by only taking into account the species of one tenthredinid tribe, namely the Phymatocerini (nine species studied). Our results support the "harmful hemolymph hypothesis" that we present here and that assumes a functional link between these morphological and chemical traits, jointly acting as a chemical defense strategy. We suspect hemolymph deterrence to be often due to sequestration of plant secondary metabolites. We discuss the role of easy bleeding, considering the fact that sawfly larvae are frequently the prey of invertebrate and vertebrate predators. It is suggested that invertebrates such as ants were more important than vertebrates in the evolution of easy bleeding.