136 Invasive Stink Bugs and Related Species (Pentatomoidea)
World and Old World species) as two separate and valid families. However, Dolling (1981) treated both
the Corimelaeninae and the Thyreocorinae as subfamilies of equal status within the Cydnidae with the
Corimelaeninae restricted to New World species and the Thyreocorinae restricted to Old World spe-
cies. Froeschner (1988e) provided an explanation for why Thyreocoridae (or -inae) had priority over
Corimelaenidae if only one family group is recognized. Štys and Davidová-Vilímová (1979) recognized
this group as a family, the Thyreocoridae. Lis (1994, 1999a) treated all New World and Old World
species as a single group, but used the name Corimelaeninae as a subfamily of the Cydnidae; he later
(2006e) recognized this group as a family (Thyreocoridae), with two subfamilies (Corimelaeninae and
Thyreocorinae). Schuh and Slater (1995) followed Dolling in recognizing this group as two subfamilies
within the Cydnidae - the Thyreocorinae for Old World taxa and Corimelaeninae for New World taxa.
The morphological analysis in the phylogenetic work by Grazia et al. (2008) supported the groupings of
Dolling (1981), but the rest of the analysis did not. They speculated that their analysis might not have been
robust enough to determine relationships or that the Cydnidae as recognized by Dolling might not be
monophyletic after all. Based on mitochondrial sequences (12S and 16S), Lis et al. (2012b) did not recover
the Thyreocoridae as a monophyletic group. Although the aim of their study was to test the monophyly
of the Dinidoridae, they included three thyreocorid species: Galgupha difficilis (Breddin), Strombosoma
impictum (Stål), and Thyreocoris scarabaeoides (L.). In the combined analysis, S. impictum and T.
scarabaeoides were defined as sister groups, thus supporting the Thyreocorinae, whereas G. difficilis
was the sister group of a clade including Acanthosomatidae, Cydnidae, Dinidoridae, Parastrachiidae,
Pentatomidae, Scutelleridae, and Tessaratomidae. Based on morphological characters from a broad
sample of included genera (12 genera of Thyreocoridae plus representatives from Canopidae, Cydnidae,
Parastrachiidae, and Plataspidae), Matesco’s (2014) study supported the monophyly of the Thyreocoridae,
including six synapomorphies: (1) dense head punctation, (2) presence of punctation between the ocel-
lus and adjacent compound eye, (3) presence of a pseudoruga (prolongation of the ostiolar ruga onto the
mesopleuron), (4) anterior margin of abdominal sternite VII angulate in males, (5) absence of a carina on
the internal surface of the dorsal rim of the pygophore, and (6) parameres partially exposed.
The relationship of the Thyreocoridae within the Pentatomoidea has been studied by several work-
ers (Gapud 1991, Grazia et al. 2008, Lis et al. 2012b, Matesco and Grazia 2015), most of whom have
indicated a close relationship with the Cydnidae. The phylogenetic study by Gapud (1991), based on
morphology, suggested that the Cydnidae and Thyreocoridae were strongly related by the presence in
both of coxal combs and tibial spines. Thyreocorids can be separated from the cydnids by the lack of
setigerous punctures on the head and thorax (present in cydnids). The Thyreocoridae shares other impor-
tant characters with some other pentatomoid families, such as the enlarged scutellum, short frenum, and
ninth laterotergites fused (Matesco and Grazia 2015). For example, the enlarged scutellum is shared with
the Canopidae, Lestoniidae, Megarididae, Plataspidae, Scutelleridae, and Pentatomidae (Aphylinae,
Cyrtocorinae, and some members of Asopinae, Pentatominae, and Podopinae); the Thyreocoridae can
be separated from all of these families by the presence of spines along the length of the tibiae.
Members of this family are relatively small (3-8 mm), oval-elongate in shape, strongly convex dor-
sally, flat ventrally, and usually a shiny black (Figure 2.25K) or dark brown color, often with some
white markings. The head is declivent in lateral view and subtriangular in dorsal view. The antennae
are five-segmented. The scutellum is greatly enlarged and strongly convex, nearly covering the entire
abdomen; the exposed part of the corium is much reduced and often of a pale whitish color. They pos-
sess a stridulatory mechanism composed of a strigil on the posterior anal vein (postcubitus in several
earlier papers) of the hind wing and a plectrum located on abdominal tergum I. In the Corimelaeninae,
the hind wings have the jugal lobe perforated (Figure 2.11F). The tibiae are provided with numerous
spines along their length, the tarsi are three-segmented, and the coxal combs are present. The abdominal
trichobothria usually are arranged transversely. The female spermatheca has a short invagination basally
(called a sclerotized funnel in Štys and Davidová 1979) that may be the precursor to a sclerotized rod; the
spermathecal bulb is simple, ball-shaped, and lacks diverticula.
The New World taxa (nine genera and 207 species at the time) were revised and keyed by McAtee and
Malloch (1933). The Old World fauna comprises three genera and six species; the European species of
Thyreocoris were revised by Štys and Davidová (1979). The Palearctic and Nearctic species recently have
been catalogued by Lis (2006e) and Froeschner (1988e), respectively. A checklist and other taxonomic