Higher Systematics of the Pentatomoidea 115
in distribution. These two groups may be related. Linnavuori (1982) believed the narrowing of the body
to be an adaptive character associated with living on narrow grass leaves. He related this group to the
Phyllocephala group based on a common prolongation of the juga, the dorsal coloration and pattern of
carinae, and the development of pale callous bands on the thoracic pleura. This tribe currently contains
three genera and 12 species (Table 2.3).
2.2.10.8.3 Phyllocephalini Amyot and Serville, 1843
Under the name Phyllocephala group, Linnavuori (1982) treated four African genera (Lamtoplax
Linnavuori, Melampodius, Nimboplax, and Phyllocephala). Kamaluddin and Ahmad (1991) transferred
the first three genera listed above to the Cressonini. Linnavuori (1982) defined this group by a number of
characters: body large, broadly ovate (Figures 2.18H, 2.27G); head shovel-shaped, with stiff hairs; of a uni-
form color; pronotum rugulose, the anterior margin with strong papillae bearing a setigerous apical pit; scu-
tellum broadly ligulate; female genital plates coarsely punctate; and the mesosternum with broad, slightly
elevated medial carinae. Again, it should be cautioned that these characters may apply to the Cressonini as
the majority of genera treated here by Linnavuori have now been transferred to the Cressonini.
The genus Delocephalus has had an interesting taxonomic history, which has involved family-level
nomenclature. Distant (1881) originally described the genus and species, D. miniatus, from Madagascar
and placed the species in the family Tessaratomidae (the spiracles on abdominal segment II were
exposed, a character thought to be important in defining the Tessaratomidae). Horváth (1900), realizing
that Delocephalus was unlike any tessaratomids he knew, erected a new division, Delocephalaria, within
the Tessaratomidae. Based primarily on its general facies, Schouteden (1909) transferred Delocephalus
to the Pentatomidae (Phyllocephalinae). Cachan (1952), apparently following Horváth, listed this genus
in the Delocephalaria within the Tessaratomidae. Kumar (1969a) studied the male genitalia and con-
firmed Schouteden’s opinion that Delocephalus was a member of the Pentatomidae. Linnavuori (1982)
included Delocephalus in his Dalsira group, but Ahmad and Kamaluddin (1990b) included Dalsira
Amyot and Serville in the Phyllocephalini. Until further work can be conducted, we tentatively treat
Delocephalus as a member of the Phyllocephalinae: Phyllocephalini, and Delocephalini is treated as a
synonym of the Phyllocephalini.
Kamaluddin and Ahmad (1988) reviewed the genera and species from India and Pakistan. This tribe
currently contains 32 genera and 167 species (Table 2.3).
2.2.10.8.4 Tetrodini Ahmad, 1981
Ahmad and Kamaluddin (1990a) defined this tribe as having the body broad and elongately ovate; the
head longer than broad and distinctly longer than the pronotum; the length anterior to the compound eyes
distinctly longer than the posterior part of head; the juga usually longer than and meeting in front of the
tylus; the antenniferous tubercles not visible from dorsal view; the humeral angles rounded, never greatly
produced; the anterolateral pronotal margins usually entire, not denticulate; and the scutellum longer
than broad. The female spermathecal bulb is ball-shaped with three relatively short diverticula, and the
duct between the sclerotized rod and the spermathecal pump is quite short (Linnavuori 1982). This tribe
currently contains four genera and 23 species (Table 2.3).
2.2.10.9 Podopinae Amyot and Serville, 1843
This subfamily, described by Amyot et Serville, 1843 (as ‘Podopides’), has not only been classified
historically within the family Pentatomidae, but specific genera (e.g., the genus Tar isa) or groups of
genera have at times also been placed within the family Scutelleridae (most likely due to the enlarged
scutellum seen in both groups). The podopines are distributed worldwide, except in the New Zealand
subregion. They represent an especially heterogenous group of species, primarily because of the diffi-
culty in classifying some of the genera/taxa (see summary of higher classification in Davidová-Vilímová
and McPherson 1995). Schouteden (1903) treated the African species of Podopinae and later (1905b) pro-
vided a summary of the Podopinae in an excellent monograph. The most recent studies concerning the