Phylogeny of the stink bug tribe Chlorocorini (Heteroptera, Pentatomidae) based on DNA and morphological data: the evolution of key phenotypic traits

ABSTRACT Pentatomidae is the third largest family of true bugs, comprising over 40 tribes. Few tribes have been studied in a phylogenetic context, and none of them have been examined using molecular data. Moreover, little is known about the evolution of key morphological characters widely used in taxonomic and phylogenetic studies at multiple levels. Here, we conduct a phylogenetic study of the tribe Chlorocorini (Pentatominae) combining 69 morphological characters and five DNA loci. We use the inferred phylogeny to reconstruct the evolution of key morphological characters such as the spined humeral angles of the pronotum, a dorsal projection on the apices of the femora and characters of male genitalia. We provide solid evidence that the tribe as currently recognized is not monophyletic based both on DNA and morphological data. The genera Arvelius Spinola and Eludocoris Thomas were consistently placed outside of the Chlorocorini, whereas the remaining genera were found to form a monophyletic group. We also show that nearly all morphological diagnostic characters for the tribe are homoplastic. The only exception is the development of the hypandrium, which contrary to expectations for genital traits, showed the slowest evolutionary rates. In contrast, the most rapidly evolving trait is the length of the ostiolar ruga, which may be attributed to selection favouring anti‐predatory behaviour and other functions of its associated scent glands. Lastly, we also provide a preliminary glimpse of the main phylogenetic relationships within the Pentatomidae, which indicates that most of the included subfamilies and tribes are not monophyletic. Our results suggest that the current subfamily‐level classification of Pentatomidae is not adequate to reflect its evolutionary history, and we urge for a more complete phylogeny of the family. We provide the first total‐evidence phylogeny within the Pentatomidae at the tribe level. According to both molecular and morphological data, the tribe Chlorocorini is not monophyletic. Nearly all diagnostic characters for the tribe are homoplastic; the only exception is the development of the hypandrium, which contrary to expectations for genital traits, showed the slowest evolutionary rates. Our results suggest that the current classification of Chlorocorini and several other tribes and subfamilies of stink bugs is not adequate to reflect the evolutionary history of the Pentatomidae.