Higher Systematics of the Pentatomoidea 111
in most phyllocephalines, reaching between the middle coxae. The mesosternum is longitudinally
carinate, the abdominal venter is unarmed, and the ostiolar rugae are relatively short, not reaching to
the middle of the metapleura.
Arrow-headed Bugs
Štys (2000), at the 21st International Congress of Entomology, reported on a “strange arrow-headed
bug” from Australia. He examined both adults and immatures that had been collected under the scales
of bark of a eucalyptus tree from South Australia and another adult collected in flight from Western
Australia. He indicated that many of its character states were shared by other Pentatominae, but it
also had many “unique” characters within the Heteroptera. Štys listed the following defining char-
acters: (1) general facies “lygaeid” or aradid-like with many morphological characters similar to the
Lygaeoidea; (2) head relatively long with lateral pre-ocular extensions; (3) bases of labrum and labium
well-separated, by at least half the length of head; (4) juga much longer than tylus, apically free in
immatures, but enclosed by anteriorly projecting bucculae in adults; (5) labium thin and long, reach-
ing nearly to genitalia; (6) reduced and somewhat unique trichobothrial pattern; (7) adults with an
L-shaped stridulatory area on each side of abdominal sternite III, and later instar immatures with
linear stridulatory areas along lateral margins of laterotergites II and III laterad of spiracles. This last
character, stridulatory areas in immatures, is only known in the Reduviidae. Štys (2000) hypothesized
that the arrow-headed bugs might belong in the Pentatominae and be related to the Diemeniini or the
Mecideini (probably because they share stridulatory areas), or that they may represent a new relict
pentatomomorphan family. This taxon still awaits formal description.
2.2.10.7.47 Fossil Genera
Placement of fossil taxa to genus, or even to tribe, can be difficult because of the condition of most fossils
and the inability to observe certain key characters. Below, we list the fossil pentatomid genera known to
us, with comments on their current classification. At least three fossil families have been described in
the Pentatomoidea. The Mesopentacoridae, and its included genera, is discussed earlier in this chapter
(see Section 2.2.8); and the relatively new fossil family, Primipentatomidae, and its included genera is
treated later (see Section 2.2.13). The third family, Spinidae, was described by Hong in 1982. The type
genus of the Spinidae, Spinus Hong, is preoccupied so Spinidae is an invalid name. Hamilton (1992),
however, placed Spinus as a junior synonym of the auchenorrhynchan genus Homopterulum Handlirsch,
effectively making the Spinidae a junior synonym of the fossil membracoid family Jascopidae.
Apodiphus robustus Jordan, 1967
Popov (2007) placed this fossil species as a junior synonym of Pentatomoides atratus (also see below).
Cacoschistus Scudder, 1890; Pentatomites Scudder, 1890; Polioschistus Scudder, 1890; Poteschistus
Scudder, 1890; Teleoschistus Scudder, 1890; Thlimmoschistus Scudder, 1890; Thnetoschistus
Scudder, 1890; and Tiroschistus Scudder, 1890
Kirkaldy (1909) asserted that all of the above listed genera may possibly be related to the carpocorine
genus Euschistus. Scudder (1890) described another fossil genus, Mataeoschistus, but this has been
placed as a junior synonym of Thnetoschistus (Carpenter 1992).
Carpocoroides Jordan, 1967; Halynoides Jordan, 1967; Pentatomoides Jordan, 1967; Pseudopalomena
Jordan, 1967; Rhomboidea Jordan, 1967; Suspectocoris Jordan, 1967
Technically speaking, these fossil generic names may not be taxonomically available. Jordan (1967)
described ten new species of fossil pentatomids, placing them into six new genera. But, he did not pro-
vide a generic description for any of the six genera, nor did he designate type species. Carpocoroides
and Pentatomoides originally included multiple species, so they are not taxonomically available (no
type species designated). The remaining four genera were originally monotypic, and so the type species
is known (by monotypy), but for these genera to be valid, ICZN Article 13.4 states that the description
must be in the form of a combined description of a new genus-group taxon and new species (if marked
by gen. nov., sp. nov., or equivalent expression). Jordan (1967) did not use this type of notation, but, in
his discussion, he states “Trotzdem habe ich 22 Arten neu benannt, vielfach sogar neue Gattungsnamen