Cladistics Cladistics 24 (2008) 1–45 10.1111/j.1096-0031.2008.00224.x Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera) Jocelia Graziaa,*, Randall T. Schuhb and Ward C. Wheelerb aCNPq Researcher, Department of Zoology, Universidade Federal do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil; bDivision of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA Accepted 18 March 2008 Abstract Phylogenetic relationships within the Pentatomoidea are investigated through the coding and analysis of character data derived from morphology and DNA sequences. In total, 135 terminal taxa were investigated, representing most of the major family groups; 84 ingroup taxa are coded for 57 characters in a morphological matrix. As many as 3500 bp of DNA data are adduced for each of 52 terminal taxa, including 44 ingroup taxa, comprising the 18S rRNA, 16S rRNA, 28S rRNA, and COI gene regions. Character data are analysed separately and in the form of a total evidence analysis. Major conclusions of the phylogenetic analysis include: the concept of Urostylididae is restricted to that of earlier authors; the Saileriolinae is raised to family rank and treated as the sister group of all Pentatomoidea exclusive of Urostylididae sensu stricto; a broadly conceived Cydnidae, as recognized by Dolling, 1981, is not supported; the placement of Thaumastellidae within the Pentatomoidea is affirmed and the taxon is recognized at family rank rather than as a subfamily of Cydnidae, although its exact phylogenetic position within the Pentatomoidea remains equivocal; the Parastrachiinae is treated as also including Dismegistus Amyot & Serville and placed within a broadly conceived Corimelaenidae, the latter group being treated at family rank; the family-group taxa Dinidoridae and Tessaratomidae probably represent a monophyletic group, but the recognition of monophyletic subgroups will benefit from additional representation in the sequence data set; and the Lestoniidae is treated as the sister group of the Acanthosomatidae. The Acanthosomatidae and Scutelleridae are consistently recovered as monophyletic. The monophyly of the Pentatomidae appears unequivocal, inclusive of the Aphylinae and Cyrtocorinae, on the basis of morphology, the latter two taxa not being represented in the molecular data set. Ó The Willi Hennig Society 2008. The recognition of a taxon with a composition similar Hypotheses of phylogenetic relationships for the to the present superfamily Pentatomoidea goes back at Pentatomoidea are presented in explicit diagrammatic least to Leach, 1815 (Leston, 1953a). Although several form in Fig. 1. These schemes, taken from Bonatto authors have considered Pentatomoidea to be a natural (1988), represent, respectively, the theories of: group, there has been substantial disagreement on the Fig. 1(a)—Singh-Pruthi, 1925 (diagram and discussion); relationships among family-level and lower categories Fig. 1(b)—Leston, 1958 (fig. 5); Fig. 1(c)—China and (Kirkaldy, 1909; Leston, 1953a; Pendergrast, 1957; Miller, 1959 (fig. 1); Fig. 1(d)—Cobben, 1968 (figs 269– Scudder, 1959; Sˇ tys, 1961; McDonald, 1966; Gross, 270); Fig. 1(e)—Cobben, 1978 (several figures and text); 1975, 1976; Sˇ tys and Kerzhner, 1975; Cobben, 1978; and Fig. 1(f)—Gapud, 1991 (fig. 28). As can be seen, Schuh, 1986; Gapud, 1991; Henry, 1997; Cassis and these classifications contain many conflicts concerning Gross, 2002). The majority of previous studies of pent- relationships among the pentatomoid families. atomoid relationships used one—or at most two—suites The first contribution on pentatomoid relationships of characters, and in many of these studies taxa were using cladistic methodology was that of Gapud (1991). He grouped on the basis of symplesiomorphic resemblance. analysed 41 characters in 13 terminal taxa, following the prior general schemes that had divided the superfamily *Corresponding author: into 11 families. Thaumastella Horva´th was included E-mail address: [email protected] within the Cydnidae; Aphylidae and Megarididae were Ó The Willi Hennig Society 2008 2 J. Grazia et al. / Cladistics 24 (2008) 1–45 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).] omitted (Fig. 1f). Although GapudÕs work was of pio- These alternative classifications of Pentatomoidea, neering importance, his matrix, and his interpretation and proposed in the last 30 years, are summarized in explanation of the characters require several modifica- Table 1. tions and corrections, which we discuss below. None of the hypotheses of relationships among taxa Schaefer (1993b) included 16 families in the Pentato- within the superfamily Pentatomoidea, as formulated by moidea. The Cyrtocoridae was treated as a family, and previous authors, is well corroborated in a rigorous following Durai (1987) and Sinclair (1989), Eumenotes cladistic context. We have therefore re-evaluated all Westwood was included in the Dinidoridae, and the available morphological evidence for 135 taxa, 84 of Oncomerinae was given family status. Sinclair (2000) which are included as ingroups in our cladistic analysis. later treated the Oncomerinae as a subfamily of the With these data we have combined as many as 3500 bp of Tessaratomidae. DNA sequence data for each of 52 terminal taxa. The Schuh and Slater (1995), in a review and synthesis of data were analysed using cladistic methods, discussed the literature, recognized 14 families within the Penta- below, in an attempt to provide a more strongly tomoidea. They treated the Cyrtocoridae as a subfamily corroborated hypothesis of pentatomoid relationships. of Pentatomidae. The ingroup includes the majority of the nominal taxa at Henry (1997), in a phylogenetic analysis of the family and below the family level, proposed to be members of groups within the infraorder Pentatomomorpha, recog- the Pentatomoidea. In the family Cydnidae, two tribes of nized five superfamilies (Pentatomoidea, Coreoidea, Cydninae (Scutellocorini Ahmad and Moizuddin and Pyrrhocoroidea, Idiostoloidea, and Lygaeoidea). Penta- Geotomini Wagner), one of Cephalocteinae (Cephalo- tomoidea was considered as monophyletic and, follow- cteini Lis), and one of Sehirinae (Amaurocorini Wagner) ing Henry and Froeschner (1988), he recognized 17 are not included. The tribe Byrsodepsini Kokorek and families within the Pentatomoidea. Henry (1997) gave Lis of the family Dinidoridae is not included. The family status to the Cyrtocoridae and Eumenotidae. recently established pentatomid subfamily Stirotarsinae Cassis and Gross (2002), in their catalogue of the (Rider, 2000) is also not included. Of the four tribes of Australian fauna, recognized within Pentatomoidea the Phyllocephalinae (Ahmad and Kamaluddin, 1988, 1990; same 14 families that received family status in Schuh Kamaluddin and Ahmad, 1988), only the nominate tribe and Slater (1995). is included. Of the five tribes of Podopinae (Davidova´- Most recently, Rider (2006) recognized 15 families Vilimova´and Sˇ tys, 1994; Davidova´-Vilimova´and McPh- within Pentatomoidea; following Sweet and Schaefer erson, 1995), three are not included. Finally, of the 42 (2002), Parastrachia Distant was given family rank. tribes of Pentatominae (Rider, 2006), 29 are not included. J. Grazia et al. / Cladistics 24 (2008) 1–45 3 Table 1 Alternative family-group classifications of the Pentatomoidea (all taxon names rendered in current spellings) Schaefer Schuh and Slater Henry and Cassis and Gross (1975) (1993b) (1995) Froeschner (1988) Gross (2002) Rider (2006) Acanthosomatidae Acanthosomatidae Acanthosomatidae Acanthosomatidae Acanthosomatidae Acanthosomatidae Aphylidae Aphylidae Aphylidae Aphylidae Canopidae Canopidae Canopidae Canopidae Canopidae Cydnidae Cydnidae Cydnidae Cydnidae Cydnidae Cydnidae Cyrtocoridae Cyrtocoridae Dinidoridae Dinidoridae Dinidoridae Dinidoridae Dinidoridae Dinidoridae Eumenotidae Lestoniidae Lestoniidae Lestoniidae Lestoniidae Lestoniidae Lestoniidae Megarididae Megarididae Megarididae Megarididae Megarididae Oncomeridae Parastrachiidae Pentatomidae Pentatomidae Pentatomidae Pentatomidae Pentatomidae Pentatomidae Phloeidae Phloeidae Phloeidae Phloeidae Phloeidae Plataspididae Plataspididae Plataspididae Plataspididae Plataspididae Plataspididae Scutelleridae Scutelleridae Scutelleridae Scutelleridae Scutelleridae Scutelleridae Tessaratomidae Tessaratomidae Tessaratomidae Tessaratomidae Tessaratomidae Tessaratomidae Thaumastellidae Thaumastellidae Thaumastellidae Thaumastellidae Thaumastellidae Thyreocoridae Thyreocoridae Urostylididae Urostylididae Urostylididae Urostylididae Urostylididae Urostylididae Monophyly of Pentatomoidea and included family-group (Reduviidae), Diplocysta sp. (Tingidae), Mezira sayi taxa Kormilev (Aradidae), Trisecus sp. (Idiostolidae), and Laryngodus sp. (Rhyparochromidae). Additional obser- Apomorphic characters supporting the monophyly of vations were made for 43 taxa listed in Table 2. Pentatomoidea, as postulated by prior authors, are: A total of 57 characters are scored; 37 of these are barrel-shaped egg structure with a circular eclosion rent binary and 20 are multistate. The multistate