60 Invasive Stink Bugs and Related Species (Pentatomoidea)
At quick glance, the edessines have a similar appearance to the rhynchocorines (Pentatominae), but
they are actually quite different. In the rhynchocorines, the base of the abdomen, the metasternum,
and the mesosternum are all produced ventrad and tightly contact each other (nearly appearing fused);
it is the mesosternum that protrudes forward over the prosternum and often onto the base of the head
(Figure 2.7E). In the edessines, only the base of the abdomen and the metasternum are produced ven-
trad; it is the metasternum that is produced forward over the mesosternum, often becoming bifid anteri-
orly over the prosternum (Figures 2.5F-H). Additionally, the rostrum is much shorter in the edessines,
and the bucculae are shorter and flap-like.
Several edessine genera have been reviewed recently, either in part or in whole: Ascra (Santos et
al. 2015), Brachystethus (Barcellos and Grazia 2003a), Doesburgedessa Fernandes (Fernandes 2010),
Grammedessa (Correia and Fernandes 2016), Lopadusa (Becker and Grazia 1970), and Paraedessa
Silva and Fernandes (Silva et al. 2013). Several species groups within Edessa have been the subject of
revisionary work (Fernandes and van Doesburg 2000a-c; Fernandes et al. 2001; Silva et al. 2004, 2006;
Fernandes and Campos 2011; Silva and Fernandes 2012; and Santos et al. 2014).
Various species may feed on a variety of plant hosts, but there seems to be some preference for mem-
bers of the plant family Solanaceae among the economically important species (Panizzi et al. 2000).
Several members of this subfamily (e.g., Lopadusa augur Stål and Edessa nigropunctata Berg) have
been reported to exhibit parental care of the young (Requena et al. 2010). There is one fossil species
described in the Edessinae, Edessa protera Poinar and Thomas (2012).
2.2.10.7 Pentatominae Leach, 1815
This is, by far, the most diverse subfamily in the Pentatomidae, containing 660 genera and 3,484 species
(Tables 2.2, 2.3). Its members occur worldwide. A taxon similar to our current Pentatominae was pro-
posed by Stål (1865) and included genera of the groups Halydes, Pentatomides, Pododides, Podopides,
Rhaphigastrides, and Sciocorides of Amyot and Serville. Later, the subfamily was divided into groups
of genera (Stål 1872, 1876). These groups eventually were rearranged and named by different authors
(Atkinson 1888, Distant 1902, Cachan 1952), and were followed partially until recently (Uhler 1886;
Kirkaldy 1909; Van Duzee 1917; Cachan 1952; Gross 1975b; Rolston and McDonald 1979; Rider 2006a,
2015a). However, depending on the characters considered important for the classification proposed, the
organization and hierarchy of the groups differ considerably (Table 2.1).
The lack of unique diagnostic characters hampers the identification of this subfamily, making
it difficult to construct a useful and stable classification. As with any large group, it is difficult to
find characters that all members possess. In general, pentatomines can be quite small (Sepontia and
Sepontiella Miyamoto species are only a few mm in length) to quite large (Catacanthus species are
nearly as large as most tessaratomids). Many of its members are dull yellows, tans, greens, browns,
and blacks, colored so as to blend in with their surroundings; others, however, may be brilliantly
colored in reds, yellows, and oranges, sometimes with a metallic sheen, perhaps aposematically
colored to advertise their repugnant odors. Most species have five-segmented antennae, but some
species only have four segments, and at least one Australian genus (Omyta) has only three segments.
The humeral angles are often simple and rounded, but, in some species, they are quite prominent
and spinuously produced. The scutellum is usually subtriangular but, in a few species, it is more
spatulate; if spatulate, it usually does not reach the apex of the abdomen. The frena usually extend
at least two-fifths the length of the scutellum (Figure 2.6F). The tarsi are usually three-segmented,
but a couple of tribes (Nealeriini, Opsitomini) and at least two genera included in other separate
tribes (Phalaecus Stål, Rolstoniellus Rider) have two-segmented tarsi.
The classification within this subfamily has been chaotic at best. The number of tribes recognized has
varied dramatically from worker to worker. For example, various workers from various parts of the world
still recognize over 40 different tribes (e.g., Cassis and Gross 2002, Derjanschi and Péricart 2005, Rider
2006a, Salini and Viraktamath 2015), and another 15-20 generic groups have been proposed (Gross
1975b, 1976; Linnavuori 1982). And yet, Schuh and Slater (1995) only recognized eight valid tribes.
There are no recent keys to all of the known tribes or genera, except on a regional basis. For example,
there are several important works covering the fauna of various Old World regions: West Palearctic