TESSARATOMIDAE Stål, 1864

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 : 967-968

TESSARATOMIDAE Stål

Historical: Sinclair (1989), in a generic revision and cladistic analysis of the exclusively Southern Hemisphere taxon Tessaratomidae , concluded that this group

is polyphyletic. He raised the Oncomerinae to family status. The Tessaratomidae under Sinclair̕s (1989) vision included two subfamilies, Tessaratominae and Natalicolinae, the latter with two tribes, Natalicolini and Prionogastrini, and the latter of those comprising two subtribes, Prionogastraria and Sepinaria. More recently, Sinclair (2000) restored Oncomerinae as a subfamily of Tessaratomidae . Rolston et al. (1993) summarized the current classification in their world

catalogue of the Tessaratomidae . Following Leston (1955), Leston (1956b) and Kumar (1969), they recognized the subfamilies Natalicolinae, Oncomerinae , and Tessaratominae, the last subdivided into three tribes:

Prionogastrini, Sepinini, and Tessaratomini. Schuh and Slater (1995) recognized the three subfamilies mentioned above, the Tessaratominae subdivided into five tribes (Eusthenini, Platytatini , Prionogastrini, Sepinini, and Tessaratomini), and the Oncomerinae into two tribes (Oncomerini and Piezosternini). Sinclair (2000) revised the Oncomerinae , including in this subfamily 15 genera, but did not recognize Leston̕s division of the subfamily into Oncomerini and Piezosternini. Monteith (2006) described maternal care of eggs and nymphs in five genera of Oncomerinae , three of them also showing nymphal phoresy where the nymphs are carried on the modified body of the female for a period after hatching.

Analytical result: Our unweighted parsimony analysis of morphological character data supports the monophyly of this group ( Figs 42 ), as does analysis under successive weighting ( Fig. 43 ), whereas analysis with PIWE under implied weights treats it as paraphyletic ( Fig. 44 ). In the combined molecular results and 52- taxon total evidence analyses under a 1: 1 cost ratio ( Figs 45 and 51 , respectively) the Tessaratomidae is always monophyletic and is the sister group of the Dinidoridae . The Tessaratomidae is paraphyletic via inclusion of the Dinidoridae in the 92-taxon combined analysis using a 1: 1 cost ratio ( Fig. 53 ), but is monophyletic using 1: 2 and 2: 2 cost ratios ( Figs 54 and 55 ). As with the Dinidoridae , a broader sample of sequence data would provide a more rigorous test of the monophyly and sister-group relationships of the Tessaratomidae and help to resolve the historical indecision as to the composition of the group.

Figures

Fig. 42. Strict consensus of 96 most parsimonious trees for full-taxon morphological data set, with unsupported nodes supressed. Length = 207; consistency index = 42; retention index = 86. (d) Non-homoplasious; (s) homoplasious.

Fig. 43. Strict consensus of three trees derived from successive weighting of the results shown in Fig. 42. (d) Non-homoplasious; (s) homoplasious.

Fig. 44. Strict consensus of 12 trees derived from implied weighting analysis of morphological data using PIWE. (d) Non-homoplasious; (s) homoplasious.

Figs 45–48. 45. One of two trees derived from analysis of combined molecular data with 1: 1 indel ⁄transition–transversion cost ratio. 46. Single tree derived from analysis of combined molecular data with 2: 2 indel ⁄ transition–transversion cost ratio. 47. Single tree derived from analysis of ~500 bp of 16S rRNA data using 1: 1 indel ⁄ transition–transversion cost ratio. 48. Single tree derived from analysis of ~1800 bp of 18S rRNA using 1: 1 indel ⁄ transition–transversion cost ratio.

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.