Higher Systematics of the Pentatomoidea 117
The spermatheca (based on several species examined; see below) has the spermathecal bulb roughly
globular in shape with 0 to 3 distal processes; the two flanges are developed, and the pumping region
between the flanges is of various patterns, and may be straight or convoluted. The sclerotized rod and
dilation are elongate, and the proximal part of the spermathecal duct is free of the rod for various lengths.
Linnavuori (1982) stated that only one pentatomine type of spermatheca (bulb small and simple) occurs
in the Podopinae, but he studied only a few species, none of which had developed processes on the bulb.
During the past several decades, several taxa have been removed from or transferred to the Podopinae.
More recently, the podopines have been accepted by most workers as a monophyletic group defined
by the following synapomorphies: hypertrophied scutellum often overlapping the apex of the abdomen
(inclusive apomorphy developed several times within the Pentatomoidea); scutellum broad/wide in the
anterior part, only slightly narrowed or even slightly wider toward the apex (large parts of the hemelytra
still uncovered by the scutellum); frena developed on the ventral surface of the scutellum, one on each
side as an oblique ledge-shaped structure reaching maximally to half of its length (autapomorphy); and
ostiolar rugae relatively small, short, and narrow. Dorsal projections of various shapes and sizes are
developed on the pygophoral area dorsad of parameres. Parandria (movable projections on the lateral
pygophoral rim) also have been mentioned as an autapomorphy of the Podopinae, but they are developed
only in some of the genera from the Podops group.
The following taxa were historically, at least for a short period, included in the Podopinae or clas-
sified as their close relatives. The genus Eipeliella originally was placed in the Graphosominen
(= Podopinae), probably due to the enlarged scutellum. Schouteden (1916) tentatively placed the genus
in the Pentatomidae near the genus Eysarcoris, which also has an enlarged scutellum. More recently,
Linnavuori (1982) placed Eipeliella in its own group and indicated that it might be related to the
Antestiini. Eipeliella has the frena located on the lateral margins of the scutellum, and the ostiolar rugae
are long and acute apically. Distant (1910a) originally placed his Australian genus Ippatha (Figure
2.24F) in the Graphosomatinae (= Podopinae), a position followed by Musgrave (1930) in his revision
of the Australian Podopinae. Gross (1975b), believing Ippatha to be distinct from the Podopinae, placed
it in its own genus group. Ippatha has an enlarged evaporative area, and the frena and dorsal processes
of the pygophore are not developed, thus distinguishing it from the Podopinae. The genus Amphidexius
never has been explicitly classified in the Podopinae, but Bergroth (1918) originally indicated a possible
relationship with the genus Dandinus, which is a member of the Podopinae. Gross (1975b) erected a
new genus group, the Ochisme group, and placed within it Amphidexius and several other related gen-
era. Amphidexius has the frena located laterally on the scutellum, and the scutellum is more triangular
in shape. The South American genus Prionotocoris originally was placed near the genus Ancyrosoma
Amyot and Serville, an Old World podopine. Grazia (1988) indicated that Prionotocoris was more
closely related to a group of genera tentatively placed in the Carpocorini, the Euschistus group. All
these genera can be separated from the podopines by the subtriangular scutellum and the lateral frena
that are relatively long, reaching two-thirds the length of the scutellum. Another South American genus,
Glyphepomis, that originally was touted as the only Neotropical member of the Podopini has been cor-
rectly moved to the pentatomine tribe Carpocorini; it differs from the Podopinae in having lateral frena
on a short, somewhat oval, scutellum that narrows somewhat apically. Finally, the tribe Procleticini
(monophyly shown by Schwertner and Grazia 2012) has been purported to be closely related to the
Podopini (Kormilev 1955), but the frena are short and lateral, and some of the genera have a subtrian-
gular scutellum.
All podopines are phytophagous, feeding on the vegetative as well as reproductive parts of plants from
different families, with groups of genera showing preferences for certain habitats and hosts (Schaefer
1981b, 1983; see below for more information). The only species considered to be pests are included in the
genus Scotinophara Stål [e.g., S. coarctata ( F.), S. lurida (Burmeister) and S. vermiculata (Vollenhoven)].
They are known to be important pests of rice in Southeastern Asia (Panizzi et al. 2000, Joshi et al. 2007).
Besides these species of Scotinophara (the most speciose genus with 40 described species), species of the
genera Podops Laporte (Figure 2.27I) and Graphosoma Laporte (Figure 2.27J) are studied the most
often (e.g., Leston 1953b; Durak and Kalender 2007, 2009; Gamberale-Stille et al. 2009).
No geographical area contains members of all the genera-groups in the Podopinae. For example, mem-
bers of the Deroploa group are restricted to the Australian region. The Nearctic region contains members