DINIDORIDAE Stål, 1867

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

DINIDORIDAE Stål

Historical: Durai (1987), in a world revision of the Dinidoridae , recognized the subfamilies Dinidorinae , with the two tribes Dinidorini and Thalmini , and

Megymeninae , also with two tribes, Megymenini and Eumenotini . Lis (1990) catalogued the Old World Dinidoridae . Rolston et al. (1996) summarized the current classification in their world catalogue of Dini-

doridae. Gapud (1991) considered Dinidoridae and Tessaratomidae to be sister groups with two synapomorphies supporting the grouping: partially exposed spiracles on the second abdominal segment (these

becoming completely exposed in Tessaratomidae ), and greatly enlarged ninth paratergites. The Dinidoridae , on the other hand, is separated from the Tessaratomidae by the reduced gonangulum and the enlarged and often reticulate hemelytral membrane, both of which characters are not found exclusively in this group. Kocorek and Lis (2000), in a cladistic revision of the Megymeninae , proposed anew tribe, Byrsodepsini , and established Eumenotini as a junior synonym of Megymenini sensu stricto.

Analytical result: After extensive searching, we were unable to secure, either through our own fieldwork or that of others, material adequate for sequencing other than for the genus Megymenum , leaving the Dinidorinae without molecular data. Our unweighted and successively weighted morphological analyses ( Figs 42 and 43 ) treat the broadly conceived Dinidoridae as paraphyletic; analysis of the morphological data under implied weights using PIWE ( Fig. 44 ) treats the Dinidoridae as a monophyletic subset of a paraphyletic Tessaratomidae . Megymenum is treated as the sister-group of Tessaratomidae in the 52-taxon molecular and combined analyses when using a 1: 1 cost ratio ( Figs 45 and 51 ). In the 92-taxon combined analyses, the Dinidoridae is paraphyletic within a larger Tessaratomidae using a 1: 1 cost ratio ( Fig. 53 ), is the monophyletic sister group of a monophyletic Tessaratomidae using a 1: 2 cost ratio ( Fig. 54 ), and also includes Urochela (Urostylididae) when applying a 2: 2 cost ratio ( Fig. 55 ). The close association of the Dinidoridae with the Tessaratomidae in our analyses is concordant with the conclusions of Gapud (1991), although there is no clearcut set of morphological characters that diagnoses that larger grouping. Future efforts should aim to produce a more broad-based taxon sample of sequence data for the group to test more rigorously its monophyly as well as its relationship with the Tessaratomidae as well as the remaining Pentatomoidea .

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.