Evolution and adaptive significance of larval midgut alkalinization in the insect superorder Mecopterida

The phylogenetic distribution of reported midgut pH values among larval Mecopterida supports a model in which the common ancestor of this group possessed an alkaline midgut, with subsequent loss of this trait in the lineage leading to the muscomorphan Diptera. The relationship between midgut pH and diet guild rank within the Lepidoptera and Diptera was tested by assigning numerical values to diet guilds (i.e., fruit, grasses, herbs, trees and shrubs, and organic detritus). Lepidopteran superfamilies were found to differ significantly in both midgut pH and in diet guild rank. Regression of mean superfamily midgut pH against mean superfamily diet guild rank yielded an R-2 of 0.79 (N = 10), whereas regression of species midgut pH against species diet guild rank yielded an R-2 of only 0.15 (N = 60). Species feeding on foliage of plant taxa high in tannins and on Solanaceae have midgut pH values above 9, and midgut pH in species feeding on these taxa is positively related to diet guild. In contrast, species feeding on the foliage of plant taxa containing terpenes, DIMBOA, glucosinolates, and pyrrolizidine alkaloids have midgut pH values near 8, and midgut pH of these species is either not related to diet guild (all species) or is negatively related to diet guild rank when the analysis is limited to the Noctuoidea. The data suggest that decreased midgut pH in species feeding on plants containing terpenes, DIMBOA, glucosinolates, and pyrrolizidine alkaloids may be an adaptive response that overrides selection for high pH in the presence of tannins and that midgut pH may be one factor contributing to the limitation of the host plant range of many species of lepidopteran herbivores.