SCUTELLERIDAE Leach

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

SCUTELLERIDAE Leach

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. 12 ). 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. 1f ). 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– 55 ) and in the analyses under 1: 1 cost ratios always basal to the Acanthosomatidae + Pentatomidae ( Figs 51 and 53 ). An improved sample of DNA sequence data across the range of scutellerid subfamilies might help to resolve this ambiguity.

Figures

Figs 7–13. 7a. Antennal segments of Megymenum sp.; 7b. antennal segments of Eumenotes obscura; 7c. flattened 2nd antennal segment of E. obscura in cross-section; 7d. antennal segments of Natalicola pallidens; 7e. flattened 2nd antennal segment of N. pallidens in cross-section; 7f. antennal segments of Phloea corticata. 8. Thaumastella namaquensis (Thaumastellidae): lygaeid type of head (Štys, 1964a); humeral angles not developed. 9. Lestonia haustorifera (Lestoniidae). Scutellum long almost attaining apex of abdomen but not covering conexivum and corium of hemelytra; base of corium expanded. 10. Canopus caesus (Canopidae), dorsal view. Well-developed scutellum, completely covering abdominal dorsum and hemelytra; spheroid form. 11. Body foliations: 11a. Phloea corticata (Phloeidae), female; 11b. Serbana borneensis (Phloeidae), male. 12. Garsauriella haglundi (Cydnidae). Corium subdivided by a medial longitudinal fracture (mlf = median longitudinal fracture). 13. Tessaratoma papillosa (Tessaratomidae). Hind wing stridulitrum. (S = stridulitrum)

Fig. 1. Diagrams showing hypotheses of Pentatomoidea classifications (Bonatto, 1988): (a) Singh-Pruthi, 1925; diagram and discussion; (b) Leston, 1958; Fig. 5; (c) China and Miller, 1959; Fig. 1; (d) Cobben, 1968; figs 269–270; (e) Cobben, 1978; several figures and text; (f) proposed phylogeny of Pentatomoidea (Gapud, 1991). [Captions removed; all taxon names rendered in current spellings; part (f) not from Bonatto (1988).]

Figs 49–52. 49. Single tree derived from analysis of ~470 bp of 28S rRNA using 1: 1 indel ⁄ transition–transversion cost ratio. 50. Single tree derived from analysis of ~1100 bp of COI mtDNA using 1: 1 indel ⁄transition–transversion cost ratio. 51. Total evidence analysis with POY of 52- taxon data set using 1: 1 indel ⁄ transition–transversion cost ratio. 52. Total evidence analysis with POY of 52-taxon data set using 2: 2 indel ⁄ transition–transversion cost ratio.

Fig. 53. One of six trees from total evidence analysis with POY of 92-taxon data set using 1: 1 indel ⁄ transition–transversion cost ratio. (d) Non-homoplasious; (s) homoplasious.

Fig. 54. One of four trees from total evidence analysis with POY of 92-taxon data set using 1: 2 indel ⁄ transition–transversion cost ratio. (d) Non-homoplasious; (s) homoplasious.

Fig. 55. One of three trees from total evidence analysis with POY of 92-taxon data set using 2: 2 indel ⁄transition–transversion cost ratio, which had the lowest MRI value. Bremer support values are shown.