Higher Systematics of the Pentatomoidea 125
foliate. Gapud (1991) removed Stirotarsus from the Asopinae and placed it tentatively in the Dinidoridae,
probably based on the roughened, greyish dorsum, similar to members of the Megymeninae. Finally, as
alluded to earlier, Rider (2000), based heavily on the presence of a dilation and sclerotized rod in the
spermatheca, returned this genus to the Pentatomidae, but he felt the remaining characters were so novel
as to warrant the erection of a new subfamily.
No biological information is available.
2.2.11 Phloeidae Amyot and Serville, 1843
This family contains three extant species in two genera (Phloeophana Kirkaldy [Figure 2.16F] and
Phloea Lepeletier and Serville [Figure 2.25G]) and one fossil species and genus (Table 2.2). The pres-
ent day species are endemic to Brazil, South America, with the core distribution in the Atlantic Rain
Forest. The single fossil species, Palaeophloea monstrosa (Heer), was described based on a fragmen-
tary impression from the Miocene of Croatia; its placement is doubtful. At times, this family has been
treated as a subfamily within the Pentatomidae (e.g., Leston 1953c), and certain workers have included
the Bornean genus Serbana within this group (e.g., Grazia et al. 2008). The phloeids lack the dilation
and sclerotized rod in the female spermathecal duct; thus, we consider them to be a family separate from
the Pentatomidae. On the other hand, members of the genus Serbana have the dilation and sclerotized
rod and, so, are treated herein as a subfamily within the Pentatomidae. The two groups have a highly
similar appearance though, being rather large (Phloeidae: 20-30 mm, Serbana: about 15 mm), ovate, and
distinctively flattened with the lateral margins of the head, pronotum, and abdomen laminate or foliate;
their mottled coloration allows them to easy blend in with the bark of trees upon which they occur. The
taxonomic position of the Phloeidae within the Pentatomoidea still is undetermined (Grazia et al. 2008).
The phloeids are characterized (Leston 1953c, Lent and Jurberg 1965, Schuh and Slater 1995) by
having the jugal foliations quite large, concealing or nearly concealing the three-segmented antennae
(Figure 2.1A); antennae are three-segmented due to anarthrogenesis (incomplete division) between seg-
ments III and IV, segment I is rather long, and segments II and III are somewhat curved. Each compound
eye is divided into dorsal and ventral sections (an apomorphy also present in Serbana, but probably
acquired independently). The scutellum is subtriangular in shape, provided with a tongue-like apical
lobe that is short in Phloea (Figure 2.25G), but greatly elongate, approaching the apex of the fore-
wing in Phloeophana (Figure 2.16F). The hemelytral membrane has a reticulate venation, and the hind
wings possess a hamus. The tarsi are three-segmented. The scent gland openings are located near the
lateral margins of the metapleura, the ostiolar rugae are rudimentary, and the vestibular scar is present.
Abdominal sternites III through VII, on each side, possess a pair of longitudinally arranged trichoboth-
ria that are located mesad of the spiracular line. In the female genital plates, paratergites IX are quite
elongate. The egg lacks a pseudoperculum.
As mentioned previously, phloeids are well suited for living on the bark of trees; their mottled dorsal
coloration looks much like lichens growing on the tree surface (Lent and Jurberg 1966). When disturbed,
but also without any apparent disturbance, they are able to discharge a defensive liquid to a considerable
distance from their anus. They also exhibit maternal care (Hussey 1934); the females protect the eggs
and at least the first instars and possibly through the first three instars. The young nymphs can actually
attach themselves to the venter of the female, and she then carries them with her (Lent and Jurberg 1965,
Guilbert 2003). The life cycle occurs entirely on the trunks of their host (Salomão and Vasconcellos-
Neto 2010, Salomão et al. 2012) with both nymphs and adults feeding on the vascular system (Bernardes
et al. 2005). Host plants include members of the plant families Euphorbiaceae, Fabaceae, Mimosaceae,
Moraceae, Myrtaceae, Rosaceae, and Urticaceae (Salomão et al. 2012).
2.2.12 Plataspidae Dallas, 1851
The family Plataspidae contains 66 genera and 606 species (Table 2.2), all restricted to the Old World
(except for two species recently introduced into the New World). Members of this group are small to
medium in size (2 to 20 mm), usually orbiculate, but occasionally more ovate or even parallel-sided;
they usually are colored in blacks and browns, sometimes with pale stripes or spots, and often somewhat