116 Invasive Stink Bugs and Related Species (Pentatomoidea)
higher classification of the Podopinae, including characters of the male external genitalia, were reported
by Schaefer (1981b, 1983). The southern European taxa were treated by Péricart (2010).
Three main trends exist in the recent higher classification of the Podopinae (again see details in
Davidová-Vilímová and McPherson 1995): (1) to demote the subfamily to the level of tribe (McDonald
1966, Ahmad 1977); (2) to view the subfamily as a monophyletic group resulting in its being divided
into two or three taxa classified as tribes or generic groups (Gross 1975b, Gapud 1991); or (3) to view
the subfamily as a monophyletic group resulting in its being divided into two to five tribes (Schaefer
1981b, 1983; Hasan and Kitching 1993; Davidová-Vilímová and Štys 1994 - but note that the two new
tribes, Deroploini and Brachycerocorini, erected by Davidová-Vilímová and Štys (1994) are nomina
nuda). Recently, Gapon (2008a) accepted only two sister groups in the monophyletic Podopinae: the
Graphosomatini sensu lato and the Podopini. The latter group was divided by Gapon into three sub-
tribes, two of which are new: the Podopina, the Kayesiina, and the Scotinopharina. Gapon (2008a),
however, did not study all taxa in these groups, and so further study is needed before these categories
are accepted conclusively.
The supposed ground plan of the ancestral podopine is as follows: The antennae are five-segmented,
with two flagellar segments longer and more slender than the other segments. The labium extends to
the middle or hind coxae, with segment I not extending beyond the posterior margins of the bucculae.
The bucculae are relatively long, simply shaped; and the ocelli are relatively small and not protruding. The
antenniferous tubercles are of the common pentatomoid pattern, located on the ventral surface of the
head close to the anteroventral margin of the compound eyes. The compound eyes are large but not dis-
tinctly protruding beyond the outline of the head. The juga are relatively simple, straight and flat, without
conspicuous structures, and are identically shaped in both sexes. The outline of the head anterior to the
compound eyes is broadly triangular without distinct hairs or bristles and/or coatings/crust.
The pronotum is relatively simple, without a longitudinal ridge or keel and lacking distinctive chae-
totaxy and/or coatings, but with a granulated area on each side anteriorly. The anterior angles of the
pronotum are rounded, without processes; the lateral angles also are rounded, not protruding; and the
anterolateral margins are simple and rounded. The scutellum lacks conspicuous structures or distinct
hairs or coatings. The frena are developed as oblique structures on the ventral surface of the scutellar
margins, reaching at most one-half of the scutellar length (Figure 2.6H). The apex of the scutellum
reaches at least to the middle of abdominal tergite VI; it is relatively broad basally, only slightly nar-
rowed apically, usually leaving almost all of the corium, most of the clavus, and the apex of the mem-
brane uncovered.
The venter of the thorax has simple episterna, without conspicuous structures. The metapleural scent
gland ostiole is accompanied by a small, short, and narrow ruga; and a relatively large, finely granulated,
flat evaporative area. The abdominal venter is unarmed at the base and usually flat to slightly convex;
and the trichobothria are paired on each side of abdominal sternites III through VII, and their alveoli
are not differentiated from the surrounding area. The posterolateral angles of the connexiva are simple
and rounded.
The pygophore (terminology after Schaefer 1977 and Davidová-Vilímová and McPherson 1991) has
its external opening directed posteriorly. The ventral wall is smooth, simple, and lacks conspicuous
structures. The ventral rim is compact; the shape ranges from nearly straight to slightly concave and
may or may not have an incision and/or process. The infolding of the ventral pygophoral rim is paral-
lel to the ventral wall and has a cup-like sclerite on the internal ventral wall that is located deep within
the pygophore. The dorsal rim and its infolding lack setae, and the rim lacks processes; the infolding
is simple, forming an obtuse angle to the dorsal wall. The lateral rim is simple, narrow, and sharp; its
outline is rounded and lacks setae and processes. The infolding of the lateral pygophoral rim, which
forms the pygophoral area, lacks setae and is simple with the dorsal processes of various shapes and
sizes. The paramere has a single arm, basic in shape, roughly simple cylindrical, with fine chaetotaxy;
a larger tubercle is present on the posterior margin, which lacks membranous structures. The phallus
(terminology adopted from Gapon and Konstantinov 2006) has a phallotheca without a shield but with
basal tubercles ventrally. The conjunctiva is larger, membranous, and has a larger number of different
membranous processes. Only the paired medial penial lobes are not fused; they are developed as sclero-
tized structures in the conjunctiva. The vesica is straight and short.