Higher Systematics of the Pentatomoidea 77
of the Carpocorini. Aulacetrus is an objective synonym of Holcogaster Fieber (Figure 2.20B); that is,
both Aulacetrus and Holcogaster have the same type species, Pentatoma fibulatus Germar. Aulacetrini
was considered valid for quite some time but now has been synonymized under the Carpocorini
(but see later discussion in this section). Oshanin (1906) also proposed a separate tribe for Holcogaster,
calling it the Holcogastraria. So, the two names, Carpocorini and Rubiconiini (and even the Aulacetrini
if Holcogaster is determined to belong in the Carpocorini) would compete for priority. The first reviser
(Atkinson 1888) used the name Carpocoraria, which now seems to be in prevailing use. Additionally, Yang
(1962) proposed the tribal name Dolycorini, based on the genus Dolycoris (Figure 2.19L), but all known
workers consider Carpocoris and Dolycoris to be closely related, definitely belonging to the same tribe.
This is the largest tribe in the Pentatominae, containing 127 genera and 503 species (Table 2.3). Its
members occur worldwide. As would be expected in a large taxon, there is considerable variation in
structure and color in this group. In general, genera of this tribe are yellowish to brownish (but other
colors including metallic greens and blues are also known), sometimes mottled, and they lack a spine or
tubercle at the base of the abdomen. The juga and tylus are usually subequal in length; occasionally the
juga are a little longer but usually not meeting in front of the tylus. The prosternum is usually flat to shal-
lowly concave (in Holcostethus Fieber [Figure 2.20C] and Peribalus Mulsant and Rey, the prosternum
is inflated, forming an obtuse ridge on either side of the rostral canal), the mesosternum is not sulcate and
usually has a well-defined medial carina, and the metasternum is usually flat to shallowly concave. The
scutellum is usually triangular in shape, occasionally more spatulate (e.g., Coenus Dallas).
According to Gross (1976), the most striking characteristic of this group is “the strong and thick
F-shaped clasper with sculptured lateral surfaces,” but even this can vary. He goes on to describe the
internal male genitalia: “The structure of the aedeagus is somewhat variable but there is a strong ten-
dency for the vesica to be robust, often sclerotized, and sometimes quite long, if the latter then not infre-
quently reflexed, curved or wound in a spiral (see McDonald re cit of Euschistus or my figure of Notius).
The medial penial plates are not infrequently absent.” The spermathecal bulb in the female tends to be
simple, ball-shaped, and without diverticula; the sclerotized rod may or may not be swollen near its apex.
Most of the New World genera presently considered to be in this tribe may not actually belong here.
There seems to be a fundamental difference in the shape and length of the ostiolar rugae between the
two geographical areas. In the Old World, the ostiolar rugae tend to be somewhat longer (still usually less
than half the width of the metapleuron), and angulate apically, with the apex attached to or detached from
the metapleuron (Figure 2.9E). In most New World genera, the ostiolar rugae are much shorter, more
auriculate in form, and the apex is often detached from the metapleuron (Figure 2.8D). The exceptions
are Antheminia, which has a couple species in the New World, but its members are distinctly related to
Carpocoris and has the Old World ostiolar ruga; and the genus Trichopepla (Figure 2.20G), which may
also be more closely related to the Old World carpocorines than those presently placed there from the
New World. There is a group of New World genera related to the well-known genus Euschistus (Figure
2.20A) that have distinctly auriculate ostiolar rugae, and, in the male genitalia, the penisfilum is dis-
tinctly elongate and coiled. It is possible that these genera will be found to be a monophyletic group and
warrant a tribe of their own.
It is impossible to find clear characters that universally separate the Carpocorini from the Pentatomini.
The structure of the thoracic venter is roughly the same in both tribes. The rugae associated with the
metathoracic scent gland ostioles are usually relatively short in the Carpocorini, but more or less short-
ened, sometimes very strongly reduced, rugae occur in several genera that otherwise clearly belong in
the Pentatomini. The base of the abdominal venter is generally unarmed in the Carpocorini, and it can be
unarmed or armed with a tubercle or spine of various lengths in the Pentatomini; this character is occa-
sionally variable even within a single genus (e.g., Pentatoma Olivier) or among closely related genera.
As a consequence, several genera (particularly those that have reduced rugae and an unarmed abdominal
venter) are difficult to place into either of these two tribes.
The Carpocorini presently contains several genera that exhibit some variable characters that may
preclude them from belonging in this tribe. Some of those are noted here. As mentioned above, Mulsant
and Rey (1866) proposed the tribe Aulacetrini for the single genus, Aulacetrus, which is an objective
synonym of Holcogaster. Cachan (1952) noted a similarity of this genus with members of the Antestiini.
Several modern workers (Stichel 1961, Putshkov 1965, Wagner 1966) treated Holcogaster as a member