130 Invasive Stink Bugs and Related Species (Pentatomoidea)
even three-segmented. The bucculae are parallel and not strongly elevated; the rostrum reaches at least to
the mesocoxae. The most obvious uniting character is the enlarged scutellum that, in most species, nearly
covers the entire abdominal dorsum; the exocorium and clavus are minimally exposed, and the frena are
short or absent; the coria are not strongly sclerotized. The hemelytral membrane has ten or more longi-
tudinal veins, and the hind wing has a hamus. In members of some scutellerid subgroups (Scutellerinae),
a strigil is found on the posterior anal vein (postcubitus in several earlier papers) of the hind wing and
a plectrum is found on abdominal tergum I. The propleura are carinate, and the prosternum is weakly
to moderately sulcate. The second gonocoxae are fused, and the first and second gonapophyses are
membranous. The gynatrium has a sclerotized basal groove, the female spermatheca is diverse, the
spermathecal duct is with or without a dilation, if the dilation is present, its proximal and distal orifices
may be provided with sclerotized tubular projections; the spermathecal bulb is simple, rounded, without
tubular diverticula. The phallus usually has two or three pairs of well-developed conjunctival projec-
tions. Twenty-seven species of scutellerids have been karyotyped, all of which have a diploid number of
10 + XY (Ueshima 1979, Kerzhner et al. 2004, Rebagliati et al. 2005).
There are other families that have an enlarged scutellum (Canopidae, Megarididae, Lestoniidae,
Plataspidae, Podopinae in the Pentatomidae, Thyreocoridae), but there are other characters that
separate this family from those listed. For example, the scutellerids lack spines and bristles on the
tibiae which is characteristic of the Thyreocoridae; the small or obsolete frena and the lack of a
pentatomid-type dilation and sclerotized rod will separate scutellerids from the Podopinae and other
Pentatomidae; the three-segmented tarsi will separate them from the Megarididae and Plataspidae,
which have two-segmented tarsi; the jugal and pronotal margins are never laminately produced as in
the Lestoniidae; and the characters of the forewing and spermatheca will separate scutellerids from
the Canopidae.
The classification within the Scutelleridae has not been stable. Schuh and Slater (1995) recognized
four subfamilies: Scutellerinae, Pachycorinae, Eurygastrinae, and Odontotarsinae; they treated the
Elvisurinae as a tribe within the Scutellerinae, and they briefly mentioned that McDonald and Cassis
(1984) had proposed a new subfamily, Tectocorinae, for the Australian genus, Tectocoris Hahn (Figure
2.26E). More recently, Carapezza (2009) erected a new subfamily, the Hoteinae (three genera). We fol-
low Eger et al. (2015a), and currently recognize eight subfamilies, three of which are subdivided into
two tribes each (Table 2.2). Most of the subfamilies occur primarily in the Old World (Afrotropical,
Australian, Oriental, and Palearctic regions) with a few genera or species in the Western Hemisphere.
The subfamily Pachycorinae is found exclusively in the New World except for one species that has
been introduced into Australia (Cassis and Vanags 2006, Tsai et al. 2011, Eger et al. 2015a). There are
currently 100 genera and 531 species (Table 2.2) placed in the Scutelleridae. There have been at least
one fossil genus and ten fossil species described in the Scutelleridae: one species in the Eurygastrinae
(Förster 1891); four species in the Odontotarsinae (Heer 1853, Meunier 1915) but one, Odontotarsus
archaicus Meunier, was transferred to the Cydnidae (Théobold 1937); one genus and two species in
the Pachycorinae (Heer 1853, 1865); two species in the Scutellerinae (Fujiyama 1967, Statz and Wagner
1950); and one species in the Tectocorinae (Henriksen 1922). The generic or even the subfamily place-
ment of several of the fossil taxa is doubtful.
The Scutelleridae was reviewed in an older but excellent paper by Schouteden (1904). Since that
time, there have been several regional studies on this family. For example, the North American fauna
was treated in an unpublished thesis (Lattin 1964); the fauna of Australia has been thoroughly studied
(McDonald and Cassis 1984, Cassis and Vanags 2006); the Taiwanese Scutellerinae taxa have been
reviewed recently (Tsai et al. 2011); and the African taxa were treated in an older paper by Schouteden
(1903). Parveen and Gaur (2015) presented an illustrated key to the scutellerid genera occurring in India,
and Leston (1952c) treated the Scutellerinae of Angola. The Neotropical fauna was treated in Eger et
al. (2015a), including a key to all genera recorded in the region. Barcellos et al. (2015) dealt with the
Argentinian fauna. Some of the more notable revisionary works for various scutellerid genera include
the following: Agonosoma Laporte (Paleari 1992), Calidea Laporte (Freeman 1939, 1946), Calliphara
Germar (Lyal 1979), Cantao Amyot and Serville (McDonald 1988), Deroplax Mayr (Ahmad et al. 1988,
1998), North American species of Eurygaster Laporte (Vojdani 1961), Lamprocoris Stål (Rédei and
Tsai 2016), Odontoscelis (Göllner-Scheiding 1986, 1987), Odontotarsus Laporte (Göllner-Scheiding