Higher Systematics of the Pentatomoidea 113
produced. The ostiolar rugae are reduced and auriculate. The digestive system in the Phyllocephalinae
is formed differently than in other pentatomine subfamilies and was referred to as “a kind of filter
chamber” by Goodchild (1966). The basal plates of the phallus are usually strongly sclerotized; the phal-
lotheca has a pair of short ventral processes that are flattened dorsoventrally, and widened posteriorly;
the ventral lobe of the conjunctiva is sclerotized and projected dorsad on each side as a spoon-shaped
process; and the vesica is short, never surpassing the ventral lobe of the conjunctiva (Linnavuori 1982,
Hasan and Kitching 1993, Konstantinov and Gapon 2005). The female spermathecal duct has a dilation
and sclerotized rod that is somewhat swollen at the base; the spermathecal bulb is ball-shaped with two
short tubular diverticula (Gross 1976).
There has been little work on the higher classification within this subfamily until recently. Linnavuori
(1982) presented an extensive discussion of the phylogeny of African genera, but refrained from using
actual tribal names. Ahmad (1981) and Ahmad and Kamaluddin (1976; 1978b; 1988; 1990a,b; 1992;
1994), in a series of papers, studied several phyllocephaline genera and erected several tribal names
to contain the Indian genera. Their classification does not totally agree with that of Linnavuori, but,
between the two, and with some intuition, most phyllocephaline genera can be placed to tribe.
Linnavuori (1982) placed the African genera in three generic groups (Table 2.6) but indicated that
there were still three genera that seemed to represent their own independent phylogenetic lineages. At the
time of Linnavuori’s work, only one tribal name was available, Delocephalini. Actually Delocephalini
was erected by Horváth in 1900 as a tribe in the family Tessaratomidae, but Schouteden (1909) (and
confirmed by Kumar 1969a) transferred the type genus, Delocephalus Distant, to the Phyllocephalinae
of the Pentatomidae without commenting on the tribal situation.
Ahmad and Kamaluddin (1976; 1978b; 1988; 1990a,b; 1992; 1994) worked extensively on the higher
classification of the subfamily, erecting three new tribal names, and assigned a number of genera to the
nominate tribe (Table 2.6). Basically, their classification goes something like this: Those genera that have
the humeral angles greatly expanded and produced anteriorly with the anterolateral margins toothed or
crenulate belong in the Cressonini; the elongate slender species belong in the Megarrhamphini; the spe-
cies that have the lateral margins of the head expanded and laminate belong in the Tetrodini; and all others
would belong in the Phyllocephalini. By combining the above classification of Ahmad and Kamaluddin
with that of Linnavuori, we have arrived at an integrated classification for this chapter (Table 2.6).
There have been few systematic works on phyllocephaline groups since the 1980s (Linnavuori 1982,
Ahmad and Kamaluddin 1988, Kamaluddin and Ahmad 1988) or early 1990s (Ahmad and Kamaluddin
1990a,b; 1992). Kamaluddin and Ahmad (1991) discussed the taxonomic position of three cressoniine
genera, Kafubu Schouteden, Melampodius Distant, and Nimboplax Linnavuori. Rider and Zheng (2004)
provided nomenclatural notes and a checklist of the Chinese species. And Arnold (2011b) described two
new species of Uddmania, bringing the total known species in this genus to three.
According to Gross (1976), “there is a strong tendency for members of this group to occupy desert or
very dry areas though a number do live in much moister, though always tropical, areas.” There are host
plant records for a handful of species; the records are predominantly from various grasses (Poaceae),
including a couple of records from bamboo. Linnavuori (1982) recorded an undetermined species of
Phyllocephala Laporte as occurring on or near the ground at the base of the grass Panicum turgidum
tussocks. He noted that most other phyllocephalines usually occur higher up on the plants.
2.2.10.8.1 Cressonini Kamaluddin and Ahmad, 1991
This tribe was proposed to contain those phyllocephaline genera that have the humeral angles greatly
expanded and produced and have the anterolateral pronotal margins toothed or crenulate. Whether these
are good phylogenetic characters remains to be determined. This tribe currently contains six genera and
12 species (Table 2.3).
2.2.10.8.2 Megarrhamphini Ahmad, 1981
This tribe has traditionally been characterized as including those genera whose species are more elongate
and slender (Figure 2.27H). Linnavuori (1982) described his Lobopeltista group in a similar fashion
although none of the genera that he discussed in this group is currently included in the Megarrhamphini.
But he dealt primarily with the African fauna whereas Megarrhamphus, and relatives, are more Asian