Higher Systematics of the Pentatomoidea 29
2.2 Pentatomoidea
As with many other superfamilies, heteropteran or not, the Pentatomoidea has a long and compli-
cated taxonomic history. Through time, each of the included families, subfamilies, and tribes has
been transferred across the family level ranks as often as there have been heteropterists working
on the classification of these taxa (see Table 2.1). Although the recognition of the most important
families (and subfamilies) has more or less stabilized, their evolutionary relationships are still poorly
understood; therefore, the current classification is certainly tentative even at the family and subfam-
ily levels. The tribal classification of most families, and particularly within the Pentatomidae remains
chaotic at best.
The superfamily Pentatomoidea is a member of the insect order Hemiptera, suborder Heteroptera,
and infraorder Pentatomomorpha. Typically, this infraorder has been divided into six superfamilies:
Aradoidea, Idiostoloidea, Pentatomoidea, Lygaeoidea, Pyrrhocoroidea, and Coreoidea. The Aradoidea
lack the trichobothria that are present in the other superfamilies; as such, those five superfamilies are
grouped together as the Trichophora, a group first proposed by Tullgren (1918). Most workers since
that time have supported the monophyly of the Trichophora (Schuh and Slater 1995). Although it was
not a focus of their study, the phylogenetic analysis of Grazia et al. (2008) also generally supported the
Trichophora as a monophyletic group. Their study also supported the monophyly of the Pentatomoidea
based primarily on four characters: (1) the enlarged scutellum, the apex of which usually reaches or
surpasses an imaginary line uniting the posterolateral angles of abdominal connexiva III, (2) the claval
commissure that is usually obsolete (i.e, the clavi usually do not surpass the apex of the scutellum),
(3) the trichobothria (usually paired but sometimes singular) located laterally near the spiracular line
on abdominal sternites II through VII, and (4) tergite VIII covering tergite IX in females. Additionally,
the five-segmented antennae may be a significant synapomorphy. Several other recent studies based on
molecular markers, including Grazia et al. (2008), also support the monophyly of the Pentatomoidea
and most of the included families. Relationships among families, subfamilies, and tribes within the
Pentatomoidea are mostly unsettled, and the available information will be discussed under each corre-
sponding section. The pentatomoid family and subfamily classification used in this chapter is outlined in
Table 2.2. We also are developing a catalog of the Pentatomoidea of the World; this work has led directly
to a series of nomenclatural changes throughout the superfamily (Rider and Rolston 1995; Rider 1998a,b,
2007; Rider and Fischer 1998; Rider and Kment 2015). At present, there are 1410 genera and 8042 spe-
cies described in the Pentatomoidea (Table 2.2). A tentative list of pentatomoid genera, and their place-
ment to family, subfamily, and tribe can be found on the internet (Rider 2015b).
2.2.1 Key to the Families of Pentatomoidea
1 Large to medium-sized species with body strongly dorsoventrally flattened; lateral mar-
gins of juga, pronotum, and base of coria greatly enlarged, foliaceous (Figures 2.1A,
2.16F); each compound eye divided into a dorsal and a ventral section by foliaceous
lateral margin of head, cryptic, bark-dwelling species (Figures 2.25G, 2.27K) ............ 2
1’ Size and shape variable, usually not conspicuously flattened; lateral margins of juga,
pronotum, and base of coria not foliaceous; if body somewhat flattened, then compound
eyes not divided into two sections .................................................................................... 3
2(1) Antennae 3-segmented (Figure 2.1A); spermatheca lacking a sclerotized rod; South
America (Figures 2.16F, 2.25G) .......................................................................Phloeidae
2’ Antennae 4-segmented; spermatheca with a sclerotized rod; Southeast Asia (Borneo)
(Figure 2.27K) ...............................................................Pentatomidae (part: Serbaninae)
3(1) Body tortoise-shaped, greatly convex dorsally, flat ventrally; margins of head, lateral
margins of pronotum, part of costal margins of coria, and lateral margins of abdomen
laminately produced ventrally (Figure 2.25D); abdominal venter of female with one or
two pairs of disc-shaped organs; Australia .....................................................Lestoniidae