Phylogeny of the South Asian Halyine Stink Bugs (Hemiptera: Pentatomidae: Halyini) Based on Morphological Characters

A phylogenetic analysis of 31 Oriental and Palaearctic genera of the tribe Halyini of South and Central Asia is presented here, concentrating upon the 22 genera for which complete data were available. Sixty-one morphological characters were analyzed using parsimony, including characters of the scent-gland apparatus and internal and external male and female genitalia. Using Phricodus as an outgroup, a single most-parsimonious tree resulted, following character reweighting according to their fit to the 52 multiple shortest trees initially found. Carenoplistus was always basal, and Jugalpada and Mustha were plesiomorphic in most bootstrapped trees. Two relatively plesiomorphic clades were identified, the Halys group (Salixocoris, Halys, and Neohalys) and the Paranevisanus group (Erthesina, Apodiphus, and Paranevisanus). Although strong bootstrap support existed for a clade of 12 genera (Tipulparra, Sarju, Izharocoris, Dalpada, Tachengia, Meridindia, Cahara, Ameridalpa, Eupaleopada, Meridalpa, Lodosocoris, and Neolodosocoris), the only consistent clades within it were a Dalpada group (Sarju, Izharocoris, Tachengia, and Dalpada) and a Lodosocoris group (Lodosocoris and Neolodosocoris). Tentative placements of the other nine genera are made, and a generic key is provided.