Grazia, Jocelia, Schuh, Randall T. & Wheeler, Ward C., 2008, Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera), Cladistics 24, pp. 932-976: 968-969

publication ID 10.1111/j.1096-0031.2008.00224.x

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Historical: This widely distributed taxon was first recognized as a family group by Leach (1815), Fieber (1861), and Stål (1867). Kirkaldy (1909) maintained the

group as a subfamily of Pentatomidae  with five tribes: Odontotarsini, Tetyrini, Scutellerini, Sphaerocorini, and Elvisurini. Van Duzee (1917) restored the group to family status. McDonald and Cassis (1984) erected a new subfamily, the Tectocorinae  , andaccepted the Elvisurinae  asavalidsubfamily. Intheirmorerecentsummaries of the

literature, Schuh and Slater (1995) and Rider (2006), following Leston (1953a), recognized four subfamilies, Eurygastrinae  , Odontotarsinae  , Pachycorinae  , and Scutellerinae  , the latter author subdividing Scutellerinae  into

three tribes: Elvisurini, Scutellerini, and Sphaerocorini. Gapud (1991) noted thatthe Scutelleridaeischaracterized  by the completely fused 2nd valvifers. Our examination reveals that the gonocoxites 9 (= 2nd valvifers) are not completely fused, asstatedby Gapud, but joinedmedially with a distinct fusion line (except in the Eurygastrinae  ) (see explanation of characters; corrections to Gapud̕s Fig. 12View Figs 7–13. 7). Fischer (2001) recognized the monophyly of Pachycorinae  , Sphaerocorinae, and Elvisurinae  , as well as a sister-group relationship between Tectocoris  and Odontotarsinae  ; he further concluded that the Scutellerinae  and Odontotarsinae  are non-monophyletic groups. Cassis and Vanags (2006) monographed the Australian genera of Scutelleridae  , updating McDonald and Cassis (1984) and Cassis and Gross (2002) in relation to the homologies and terminology of morphological characters. They also discussed the current literature on the monophyly and supra-familial position of the Scutelleridae  within the Pentatomoidea  , as well as for the infrafamilial classification of the scutellerids, recognizing five subfamilies.

Analytical result: Although the status of the Scutelleridae  has been debated ( Lattin, 1964; Kumar, 1965; McDonald and Cassis, 1984; Fischer, 2001; Cassis and Gross, 2002; Cassis and Vanags, 2006), our analyses offer support for the concept of a monophyletic taxon, one that is reinforced by the morphological, molecular, and combined analyses. In addition to the morphological data, this conclusion is based on a reasonably good sample of previously unavailable DNA sequence data for one of the six recognized subfamilies. Morphological characters supporting scutellerid monophyly include one synapomorphic character: areas surrounding orificium receptaculi, in pars communis, with an elongate, grooved sclerite (541). The exact relationship of the Scutelleridae  within the pentatomoid hierarchy is less clear, however. Gapud (1991) treated the Scutelleridae  as the sister group of the Canopidae  , the two groups having a central position in the cladogram ( Fig. 1fView Fig). Our total evidence analyses are ambiguous as to the precise placement of the Scutelleridae  , but always put the group distal to the Plataspididae  and Parastrachiidae  ( Figs 51– 55View Figs 49–52. 49View FigView FigView Fig) and in the analyses under 1: 1 cost ratios always basal to the Acanthosomatidae  + Pentatomidae  ( Figs 51View Figs 49–52. 49 and 53View Fig). An improved sample of DNA sequence data across the range of scutellerid subfamilies might help to resolve this ambiguity.