Higher Systematics of the Pentatomoidea 105
treat the genera from this group as members of the Aeschrocorini. This may or may not be valid, but see
discussions under that tribe (see Section 2.2.10.7.4).
2.2.10.7.45.2 Acoloba Group Linnavuori (1982) proposed this group to accommodate a single genus,
Acoloba (Figure 2.21D), a genus endemic to Africa. He indicated that it was probably related to the
Diploxys group (= Diploxyini) (see Section 2.2.10.7.20), but differed from that group by its greenish
coloration; the body being quite elongate, narrow, and depressed; the bucculae gradually tapering basad;
the anterolateral pronotal margins being narrowly reflexed; the external structures associated with the
metathoracic scent glands; and by the structure of the male pygophore. The ostiolar rugae are longer, and
the evaporatoria are distinct, well delimited from the shiny and punctate lateral margins of the meta-
pleura. The male pygophore is slender, apically trilobate; the male parameres are strongly curved. The
spermathecal bulb is broad and rather flat, with two tubular diverticula.
Linnavuori (1982) stated that this group displayed “several advanced characters connected with the
specialized mode of life on grass leaves.”
2.2.10.7.45.3 Aeliomorpha Group This is another monotypic group, containing only the genus
Aeliomorpha that has several species occurring in Africa, the Middle East, and into the Indian region.
Linnavuori (1982) stated that this was a well-known group and noted that many previous workers had
regarded it as a close relative of the genus Aelia, the type genus for the tribe Aeliini (see Section 2.2.10.7.2).
In fact, in some phylogenetic papers (Ahmad et al. 1990, Zaidi 1994), Aeliomorpha was used to represent
the tribe Aeliini, which of course may have given erroneous results as this genus is no longer accepted as
a member of the Aeliini. Linnavuori (1982) recognized possible relationships with the Aeliini and possibly
the genus Acoloba but concluded that they were not contribal. He listed a few characteristics that separated
Aeliomorpha from the Aeliini: the body is less convex, the juga and tylus are subequal in length, the anten-
nae are sexually dimorphic (segment II in males is relatively short), the propleura are not lobed anteriorly
over the base of the antennifers, the mesosternum is carinate (not sulcate), the parameres are shaped dif-
ferently, and the female spermathecal bulb is simple and ovate. Stål (1865) treated Aeliomorpha as a sub-
genus of the genus Pentatoma, but, at that time, his concept of Pentatoma was quite broad and contained
a number of subgenera that are now placed in the Carpocorini (see Section 2.2.10.7.12) and Pentatomini
(see Section 2.2.10.7.34). Cachan (1952) formally treated Aeliomorpha as a member of the Carpocorini.
Linnavuori (1982) stated that members of this genus are “grass feeders in various savanna habitats,
and in meadows, fields, etc.”
2.2.10.7.45.4 Carbula Group Linnavuori (1982) included seven genera in his Carbula group –
Aspavia (Figure 2.21G), Carbula (Figure 2.21H), Durmia, Empiesta Linnavuori, Gynenica, Mulungua
Linnavuori, and Pseudolerida. He devoted much discussion to the interrelationships of the included gen-
era but a much less effort to actually defining the group. He stated that the Carbula group was a uniform
group. He characterized the group as having a carinate mesosternum; ostiolar rugae that are relatively
short but with fairly large, well-delimited evaporative areas; and the male parameres that are biramous.
The female spermathecal bulb is ball-shaped, lacking diverticula.
Most workers have treated these genera as members of the tribe Eysarcorini (see Section 2.2.10.7.22),
although the scutellum is not as large as in most eysarcorine genera, with one exception – the genus
Gynenica now is regarded as a member of the Lestonocorini (see Section 2.2.10.7.25).
2.2.10.7.45.5 Veterna Group Linnavuori (1982) stated that he “studied” nine genera belonging in this
group, implying that there probably were other genera that also belonged here. The nine he studied
were Benia, Caura, Lerida, Leridella Jeannel, Lokaia, Mabusana, Paralerida, Tripanda, and Veterna
(Figure 2.19H). Most of the included genera in this group are brownish in color, have the abdominal
venter unarmed, and have the ostiolar rugae relatively long, all characters shared with the Cappaeini (see
Section 2.2.10.7.11). As such, these genera have been discussed under that tribe.
2.2.10.7.45.6 Halyomorpha Group Linnavuori (1982) included four genera in this group – Adelocus
Bergroth, Boerias (Figure 2.19G), Halyomorpha (Figure 2.28L), and Hymenomaga Karsch. All of