Higher Systematics of the Pentatomoidea 49
(e.g., Acoloba Spinola [Figure 2.21D], Aelia F. [Figure 2.28A], Mecidea Dallas [Figure 2.30F]). All
variations of coloration exist, from browns and greens allowing the individuals to blend in with their
surroundings, to bright reds, oranges, and blues, sometimes in metallic hues (Figures 2.27A-2.32L). The
family is characterized further (Schuh and Slater 1995, Grazia et al. 2015) by the antennae usually hav-
ing five segments, although some subfamilies (Cyrtocorinae, Serbaninae) and species only have four, and
at least one halyine genus, Omyta Spinola, has only three. The scutellum is usually large and subtriangu-
lar, although it may be enlarged in some groups, and even cover a large portion of the abdomen in some
asopines, podopines, and pentatomines. Frena are present, and they usually extend beyond the middle of
the scutellar margins; the clavi usually do not extend beyond the apex of the scutellum, so the claval com-
missure is lacking. In many groups, the thoracic sterna are sulcate medially, whereas in more advanced
groups, the sulcus has been replaced by a medial carina. The tarsi are usually three-segmented, but they
are only two-segmented in the Cyrtocorinae, Stirotarsinae, and in some minor pentatomine tribes or iso-
lated genera. The abdominal trichobothria are usually transverse, and located near the spiracular line. In
some species, there are areas on the abdominal venter on each side that appear to be thinner, somewhat
opaque; these “cuticular patches” were studied in a series of papers (Staddon 1992, 1998, 2000; Staddon
and Ahmad 1994; Staddon et al. 1994). The female spermathecal duct has a dilation with a long, slender,
sclerotized rod evaginated from the distal orifice; the spermathecal pump is well-developed with both a
proximal and a distal flange; the spermathecal bulb may be simple, digitoid, or ball-shaped, often with
one to three tubular diverticula. The eggs are usually barrel-shaped with a pseudoperculum. Nearly 300
species of Pentatomidae have been karyotyped; the diploid number varies from 10 + XY up to 24 + XY,
but the most common diploid number by far is 12 + XY (Ueshima 1979, Kerzhner et al. 2004, Rebagliati
et al. 2005).
Almost all subfamilies of Pentatomidae are well-defined by unique apomorphies, supporting the mono-
phyly of these taxa (Rolston and McDonald 1979, Gapud 1991, Konstantinov and Gapon 2005, Campos
and Grazia 2006, Gapon and Konstantinov 2006). The only exception is the subfamily Pentatominae,
a ‘catch-all’ taxon that includes several genera and groups of genera not recognized in any of the other
subfamilies (Cassis and Gross 2002).
The phylogenetic relationships among pentatomid lineages have been almost completely ignored.
Leston (1953a, 1954a) was the first to document and discuss the monophyly and phylogenetic relation-
ships within the Pentatomidae. McDonald (1966), Gross (1975b), and Linnavuori (1982) expanded these
studies and suggested good apomorphic characters for the recognition of several monophyletic groups
within the family. Gross (1975b) and Linnavuori (1982) also discussed possible phylogenetic relation-
ships among those groups. Gapud (1991) and Hasan and Kitching (1993) were the first authors to discuss
phylogenetic relationships with the Pentatomidae based on phylogenetic trees.
More recently, studies under a phylogenetic framework have been conducted at different taxo-
nomic levels. The monophyly of the subfamily Edessinae was tested in part by Barcellos and Grazia
(2003b); they also proposed hypotheses concerning the relationships among the included genera. The
work of Campos and Grazia (2006) supported the monophyly of the tribe Ochlerini (Discocephalinae);
Garbelotto et al. (2013) and Roell and Campos (2015) expanded the knowledge about phylogenetic rela-
tionships within this tribe. Concerning the subfamily Pentatominae, phylogenetic studies at the tribal
(Schaefer and Ahmad 1987, Memon et al. 2011, Schwertner and Grazia 2012), groups of genera (Grazia
1997, Bernardes et al. 2009), and genera levels (Thomas 1985, Fortes and Grazia 2005, Ferrari et al.
2010, Greve et al. 2013) have been published. But most of the subfamilies, tribes, and groups of genera
have never been studied in a phylogenetic framework, and the relationships within the Pentatomidae
remain unknown. The results of Gapud (1991) and Grazia et al. (2008) suggest that the Aphylinae and
Cyrtocorinae may be basal lineages, but a definitive conclusion and better resolution at the family and
tribal levels in the phylogenetic hypothesis presented were limited by the scope of the study conducted.
The pentatomid subfamilial and tribal classification used in this book is outlined in Table 2.3.
We will not go into the biology of the Pentatomidae here. Some notes on biology, habitats, etc., will be
discussed under the subfamily and tribal sections, and also in many of the chapters throughout this book.
The Australian, Iranian, North American, and Palearctic species have recently been catalogued by Cassis
and Gross (2002), Ghahari et al. (2014), Froeschner (1988c), and Rider (2006a), respectively. Salini and
Viraktamath (2015) provided keys for the identification of the South Indian genera. Grazia et al. (2015)