UROSTYLIDIDAE Dallas, 1851

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 : 963-964

UROSTYLIDIDAE Dallas , sensu stricto Historical: The systematic position of this family has been ambiguous over time, as was stressed by Schuh and Slater (1995). Singh-Pruthi (1925) related it

to the Acanthosomatidae . Yang (1938a,b, 1939) and Pendergrast (1957) related it to the Pyrrhocoridae , and Miyamoto (1961) to the Pentatomidae . Kumar (1971) believed that the group represented an early divergence from the other pentatomomorphans, possibly together with the Pyrrhocoridae , with which they share uniquely the fused 2nd valvifers that form an M- or W-shaped sclerite. As Gapud (1991) did not provide an illustration of what he considered as separate 2nd valvifers, we conclude that he erroneously interpreted the 2nd valvifers in the species of Urostylididae that he studied. Berger et al. (2001) proposed the emended spelling Urostylididae , in order to remove the homonymy between the heteropteran family name and Urostylidae Bütschli (Ciliophora, Hypotrichia).

Analytical result: The treatment of the Urostylididae as the basal grouping within the Pentatomoidea by Gapud (1991) is a position supported by our morphological analyses ( Figs 42–44 ) and combined analyses under an unweighted equal costs regime ( Figs 51 and 53 ). Under unequal costs, Urochela is dissociated from Urostylus and moved up to a more central position within the pentatomoid cladogram. Characters supporting the treatment of the Urostylididae sensu strico as the sister group of all remaining Pentatomoidea are listed below in the Conclusions. The monophyly of the Urostylididae sensu strico is

recognized in our analysis by the presence of a stridulitrum on A 1 in the hind wing (221) and the presence of bristles on the claws (311). Additionally, gonocoxites 9 (in females) form an Mor W-shaped

sclerite (465; Fig. 28 ), which we suggest argues for its treatment at the family level.

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 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.

Figs 25–34. 25. Parastrachia japonensis (Parastrachiidae), male. Sternite VIII visible, not covered by segment VII. 26. Tessaratoma papillosa (Tessaratomidae), female. Tergite IX visible dorsally. 27. Pallantia macula (Pentatomidae), female. Tergite IX not visible dorsally, covered by apically positioned tergite VIII. 28. Urochela distincta (Urostylididae), female. Gonocoxites 9 in an ‘‘M’’ sclerite. 29. Phloea subquadrata (Phloeidae), female. Gonocoxites 9 completely fused. 30. Tectocoris diophtalmus (Scutelleridae), female. Gonocoxites 9 fused, with a distinct median fusion line. 31. Similiforstona bella (Pentatomidae), female. Laterotergites 9 contiguous covering segment X. 32. Pantochlora vivida (Pentatomidae), female. Laterotergites 9 separate with segment X between them. 33. Lestonia haustorifera (Lestoniidae), female. Laterotergites 9 totally fused with segment X concealed. 34. Prionogaster serratus (Tessaratomidae), female. Laterotergites 9 larger than gonocoxites 8. g9, gonapophyses 9; gc8, gonocoxites 8; gc9, gonocoxites 9; go, gonangulum; la8, laterotergites 8; la9, laterotergites 9; M, ‘‘M’’ sclerite; R2, second rami; VII, seventh segment; X, tenth segment.